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BOOKS    BY 
WILLIAM    EMERSON     RITTER 

The  Higher  Usefulness  of  Science. 

The  Probable  Infinity  of  Nature 
AND  Life. 

The  Unity  of  the  Organism,  or 
the  Organismal  Conception  of 
Life.     Illustrated. 

The  Unity  of  the  Organic  Species, 
WITH  Special  Reference  to  the 
Human  Species. 

War,  Science  and  Civilization. 

An  Organismal  Conception  of 
Consciousness. 


RICHARD  G. badger, publisher,  boston 


FIGURE    56.       SKELETOX    OF    PYTIIOX. 


THE    UNITY    OF    THE 
ORGANISM 

OR 
THE    ORGANISMAL    CONCEPTION    OF    LIFE 


BY 

WiTXiAM  Emerson  Ritter 

Director  of  the  Scripps  hislitiitlon  fur 

liiologiiul  Research  of  the  Univirsity 

of  Calif ornia.  La  Jolla 

California 


TWO  VOLUMES 
VOLUME  TWO 


ILLUSTRATED 


BOSTON 

RICHARD  G.  BADGER 

THE    GORIIAM    PRESS 


Copyright,  1919,  by  Richard  G.  Badger 


All  Rights  Reserved 


This  work  contains  the  text  of  the  book: 
"An  Organismal  Theory  of  Consciousness." 


Made  in  the  United  States  of  America 


The  Gorham  Press,  Boston,  U.  S.  A. 


-z. 


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^CL 


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%)NTENTS  ^^%, 


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?^RT  I 

CRITIQUE   OF   THE   ELEME^^TALIST  CONCEPTION    OF   THl. 

ORGANISM 

B.     The  Production  of  Individiuils  by  Other  Iiulir'nlual.s    {<'i,iirlii(ltii) 

CHAPTER  PACK 

XIV.  Evidence    from    Metazoax    Germ-Cells  That    Substances 

Other  Than   Chromatin    Are   the   Physical   Bases   of 
Heredity  ..........         I 

Evidence  from  spermatozoa,  1.  Spermatozoa  sultjcrf  to 
heredity  as  well  as  "bearers  of  heredity,"  1:  (a)  ////<.*- 
trated  by  the  ontogeny  of  mamm^ilian  sperm,  4;  {b)  Illus- 
trated by  the  ontogeny  of  an  insect  sperm,  9.  Evidence  frt^m 
the  ovum,  15:  (a)  eggs  of  ascidians — the  facts,  HI;  (b) 
Conklin's  interpretation,  19.  Critical  examinatix)n  of  Conk- 
lin's  interpretation,  23. 

XV.  Evidence  from  Somatic  Histogenesis  in  the  Mui.ticei.i  ri.vR 

Organisms 32 

The  mitochondrial  theory  of  heredity,  32.  The  mitochon- 
drial theory  tested  by  the  ontogeny  of  spermatozoa,  .ill.  The 
mitochondrial  theory  tested  by  histogenesis.  ,/,\  The  un- 
tenable hypothesis  that  cytoplasm  of  the  ovum  /.v  inheritance 
material  for  general  but  not  for  special  characters,  /fO.  Spe- 
cies attributes  in  single  cells  of  adult  (nganisnis.  //.I.  The 
spiwules  of  the  ascidian  genus  slyela,  Jf.'f.  The  spicuhs  of 
sponges  and  other  invertebrates,  oO.  The  "hair.s"  of  hight  r 
phmts,  55.  Cell~7vall  structures  in  hightr  plants,  57.  The 
morphology  of  striated  muscU  ptx'rs,  60.  Tlw  physiology  of 
muscle  fibers,  61.  Summary  of  positive  information  tUxuif 
the  physical  bajtis  of  heredity,  6'f. 

XVI.  The    Inheritance    Materials    of    Gehm-Cei.i.s     Initiators 

Rather    Than    Determiners         ......       Ctd 

Antecedents   of   the   cytoplasmic  and   nuclear  thiories   of   i»»- 

ix 


i;j 


o'-^ 


xu  .        Contents 

CHAPTER  PAGE 

reasoning,  S06.  Theories  of  animal  behavior  in  relation  to  the 
"science"  of  zoology,  208. 

XXII.  Psychical  Integration 214« 

Preliminary  remarks:  (a)  Absolute  discrimination  between 
re/lex  and  psychical  phenomena  not  necessary,  214;  {b)  The 
organism  an  original  datum  in  all  problems  of  psychic  life, 
215;  (c)  Provisional  classification  of  psychical  facts,  217. 
Likeness  between  tropistic  and  higher  psychic  activity,  220. 
First  move  toward  shoiving  the  organi^mal  character  of  the 
higher  psychic  life,  227.  Associationist  psychology  a  special 
case    of   elementalist    biology,   228.      Preliminary    examination 

of  objective  and  subjective,  231.  The  essence  of  Wundtian 
apperception,  232.     Remarks  on  analysis   and  synthesis,  236. 

XXIII.  Organic  Connection  Between  Physical  and  Psychical    239 

A  still  closer  look  at  the  organismal  nature  of  tropisms, 
239.  The  automatic  and  anticipatory  character  of  tropisms 
and  other  reflexes,  241-  A  still  closer  look  at  the  likeness 
between  higher  rational  life  and  tropisms,  242.  A  still 
closer  description  of  the  subrational  moiety  of  psychic  life, 
246.  Remarks  on  the  classes  of  subrational  life,  246.  Four 
certainties  about  the  adaptiveness  of  subrational  psychic 
activities,  250.  Generally  useful  to  individual  and  to  species, 
250.  Many  useful  to  species  primarily,  250.  VafiabiUty  of 
subrational  activities,  251.  Tendency  of  subrational  activ- 
ities to  excessiveness,  256.  Summary  of  organisnud  charac- 
ter of  all  subrational  2}sychie  life,  274-  Specificity  of  subra- 
tional psychic  life,  276. 

XXIV.  Sketch  of  an   Organismal  Theory  of  Consciousness     .     283 
Remarks  on  the  hypothetical  character  of  this  chapter,  282. 

The  natural  history  method  and  the  study  of  one's  self,  282. 
Formulation  of  the  central  hypothesis,  286.  Preliminary 
justification  of  the  hypothesis  as  such,  287.  More  systematic 
justification  of  the  hypothesis,  291.  The  nature  of  "outer" 
or  objective  and  "innef  or  subjective,  292.  As  to  the  lowest 
terms  of  self -consciousness,  308.  Instinct  and  physical  or- 
ganization, 310.  Emotion  and  physical  organization,  316. 
Glance  at  the  equilibrative  interaction  between  "body"  and 
"soul,"  323.     Support  of  the  hypothesis  by  the  physico-chem- 


Contents  xiii 

CHAPTER  PAGE 

ical  conception  of  the  organism,  32/f.  Personality  and  ele- 
mentary chemical  sut)stances,  327.  On  the  psychology  of 
subjective  and  objective  personality,  331.  Personality  and  the 
"breath  of  life"  viewed  in  the  light  of  physical  chemistry 
of  the  organism,  336. 

Postscript         .         .         .         .         .         .         .         .         .         .         .         .351 

BiBIJOGRAPlIY  ...........     359 

Glossary  ............     377 

+^DEX  »»»«». .tt'*..      oJ I 


4 


LIST  or  ILLUSTRATIONS 

FIGURE  PAGE 

56.  Skeleton   of    Python Frontispiece 

36.  Sperm  of  Fur  Seal   (After  Oliver) 

37.  "Young"  Spermatid   (After  Ballowitz) 

38.  S])ermati(l  of  Fur  Seal  (After  Oliver) 

39.  Spermatid  of  Fur  Seal   (After  Oliver) 

40.  Spermatid  of  Fur  Seal   (After  Oliver) 

41.  Spermatid  of  Fur  Seal   (After  Oliver) 
43.  Development  of  Sperm  of  Argas  Miniatus  (After  Casteel)       10,  11 

43.  Development  of  an  Ascidian  Egg   (After  Conklin)   ...  17 

44.  Spinule  Cell  of  Styela  Yakutatensis   (After  Huntsman)           .  44 

45.  Spinule  Cell  of  Styela  Plicata  (After  Huntsman)     ...  45 

46.  Spinule  Cell  of  Styela  Greeleyi   (After  Huntsman)           .         .  46 

47.  Spinule  Cell  of  Styela  Montereyensis   (After  Rittcr  and  For- 

syth)         . 47 

48.  Spicules  of  Sponges    (After  I.ankester) 51 

49.  50,  51.     Development  of  a  Spicule    (After  Lankcster)       .         52,  53 

52.  Trichomes  of  Papaver  Orientalc   (After  Cannon)     .         .         .  55 

53.  Trichomes  of  P.  Pilosum    (After  Cannon)          ....  55 

54.  Trichomes  of  P.  Somniferum   (After  Cannon)  ....  55 

55.  Side  View  of  Amphioxus   (After  Parker  &  Haswell)       .         .  95 

57.  Tentacle  of  Halocynthia  Johnsoni    (Schematic;   After   Kilter)  98 

58.  Acacia  Elata 99 

59.  Vicia   Gigantea KM) 

60.  Cassia    Sp KM) 

61.  Sequoia   Sempervirens K)l 


XV 


PART  I 

CRITIQUE    OF    THE    ELEMENTALIST    CONCEPTION    OF 

THE    ORGANISM 

B.     The  Production  of  Individuals  by  Other  Individuals 


%. 


\ 


.^5 

THE  UNITY   OF  THE  G^^NISM 

Chapter  XIV  ^^^ 

EVIDENCE  FROM  METAZOAX  GERM-CELLS  THAT 
SUBSTANCES  OTHER  THAN  CHROMATIN 
ARE  THE  PHYSICAL  BASES  OF 
HEREDITY 

Evidence  from  Spermatozoa 

T  N  our  discussion  we  will  be  guided  by  the  principle  laid 
-*■  down  earlier  and  followed  throughout  our  treatment 
of  heredity  in  the  Protozoa,  namely,  that  descriptive  onto- 
genesis brought  to  bear  on  the  actual  transformations  which 
result  in  the  production  of  specific  organs  and  parts,  is  the 
final  tribunal  for  the  determination  of  what  substances  are 
hereditarily  formative.  The  first  inquiry  will  Ik?  whether 
there  exist  among  the  metazoa  single-cell  organs  or  parts 
which  when  fully  formed  exhibit  species  characters  in  the 
sense  of  taxonomic  biology.  If  such  elements  do  exist,  ob- 
servation on  the  constituents  of  the  undifTerentiated  cells 
which  take  part  in  the  transfonnations,  obviously  may  be 
expected  to  give  us  the  information  sought  as  to  what 
substances  are  formative. 

Spermatozoa  Subject   to  Heredity  as   Well  as  "'Bearers  of 

Heredity** 

The  comparative  anatomy  and  comparative  ontogenesis 
of  the  male  germ-cells  among  animals,  which  have  been  pur- 
sued with  great  assiduity   and  skill  during  recent  decades, 

1 


2  The  Unity  of  the  Organism 

furnish  perhaps  the  largest  mass  of  relevant  facts  we  pos- 
sess from  any  one  field. 

Innumerable  researches  on  fully  formed  spennatozoa,  the 
greatest  single  research  being  that  of  Retzius,  give  us  ex- 
tensive knowledge  of  the  variety  of  structure  of  the  sperma- 
tozoa in  the  larger  and  smaller  taxonomic  subdivisions  of 
the  animal  world.  It  would  be  going  beyond  the  evidence 
to  say  that  every  well-characterized  animal  species  may  be 
identified  by  its  speraiatozoa ;  but  unquestionably  the  trend 
of  investigation  has  been  toward  such  a  conclusion.  I 
believe,  for  example,  it  would  be  impossible  to  assert  on  the 
basis  of  evidence  that  any  two  species  of  animals  belonging 
to  different  genera,  no  matter  how  much  alike  if  their  dis- 
tinctiveness is  not  questioned,  have  indistinguishable  sper- 
matozoa. "One  may  say,"  writes  Ballowitz,  "that  each  ani- 
mal species  has  its  own  sperm-form  of  definite  size."  ^  An 
attempt  to  illustrate  fully  this  variety  of  form  and  size  by 
specific  examples  is  out  of  the  question  here.  We  will  refer 
only  to  the  specificity  of  the  sperm  of  man  himself.  Ret- 
zius was  able  to  compare  in  detail  sperms  of  the  Chimpanzee, 
Orang-Utan,  Gibbon,  and  Homo,  and  found  that  while  their 
resemblance  is  rather  close,  each  possesses  clear  differential 
marks.  For  example,  the  spiral  structure  of  the  envelop  of 
the  central  piece  is  considerably  more  distinct  in  the  Chim- 
panzee than  in  Homo.  Worthy  of  mention  is  the  fact  that, 
according  to  Retzius,  the  sperm  of  the  Chimpanzee  resembles 
that  of  Homo  more  closely  than  does  that  of  the  Orang, 
thus  falling  in  with  the  fact  that  in  several  particulars  of 
adult  structure  the  resemblance  of  the  Chimpanzee  to  man 
is  closer  than  that  of  the  Orang. 

The  spermatozoa  of  a  given  animal  group  having  a  closer 
resemblance  to  one  another  than  to  those  of  other  groups; 
in  other  words,  having  a  resemblance  due  to  descent,  are 
themselves  subject  to  heredity  and  are  not  alone  concerned 
in  the  transmission  of  hereditary  attributes  from  parent  to 


Evidence  from  Metazoan  Germ-Cells  3 

offspring. 

Especially  lm])()rtant  for  us  Is  it  to  notice  that  to  a  great 
extent  the  diversity  of  structure  among  s])erniatozoa  is  in 
the  locomotor  organ,  the  tail;  that  is,  the  organ  cliieflv 
concerned  with  the  unique  life  of  the  sj)erm  as  such,  and 
very  little  if  at  all  concerned  directly  with  fertilization  and 
hence  with  hereditary  marks  of  offspring.  This  fact  de- 
serves attentive  consideration.  "In  its  more  usual  form  tlie 
animal  spermatozoon  resembles  a  minute,  elongated  tad- 
pole, which  swims  very  activel}^  about  by  the  vibration  of  a 
long,  slender  tail."  -  In  some  respects  comparison  of  the 
spermatozoon  of  the  type  here  indicated  with  an  Ap])en- 
dicularian,  a  minute  Tunicate  which  possesses  a  tail  through- 
out its  life,  is  more  instructive.  Any  one  who  has  had  op- 
portunity to  observe  both  sperm  cell  and  appendicularian 
when  alive  and  active  will  not  have  failed  to  remark  the 
general  resemblance,  not  only  as  to  form  but  as  to  kind 
of  movement  in  the  two  cases.  Is  the  develoj^ment  of  tlie 
Appendicularian's  tail  a  manifestation  of  heredity.''  Surely 
no  one  would  think  of  giving  any  but  an  affirmative  answer. 
How,  then,  deny  that  the  development  of  the  spermatozoon's 
tail  is  also  a  manifestation  of  heredity.^  I  cannot  see  that 
it  would  be  less  inconsistent  to  affirm  that  the  wriggling  ap- 
pendicularian is  alive  but  that  the  wriggling  spermatozoon 
is  not,  than  to  affirm  that  the  ontogeny  of  the  first  is  guideil 
by  heredity  while  that  of  the  second  is  not.  TO"  bring  the 
point  onto  somewhat  more  familiar  ground,  let  us  revert 
to  Wilson's  comparison  of  the  spermatozoon  to  the  tadpole 
stage  in  the  life  of  the  frog.  Our  contention  is  that  the  tail 
of  the  frog's  spermatozoon  is  as  indisputably  modeled  by 
heredity  as  is  the  tail  of  the  frog's  tadpole,  and  consequently 
that  we  are  bound  to  search  for  the  physical  basis  of  hered- 
ity in  the  former  as  well  as  in  the  latter.  Our  inquiry  is, 
then.  What  observations  have  we  as  to  the  substances  con- 
cerned in  producing  the  spermatozoon  tail.? 


4  The  Unity  of  the  Organism 

(a)  Illustrated  hy  the  Ontogeny  of  Mammalian  Sperm 

Without  exception,  so  far  as  I  know,  positive  description 
of  spermatogenesis   affirms  that  the  tail  is   produced  from 

h-c, 


vi-:SS»i>;:i 


■CI  TX, 


ra.p. 


FIGURE    36.       SPERM    OF    FUR    SEAL    ( AFTER    OTJ^^ER). 

h.c,  head  cap.  n'k.,  neck,  c.p.,  connecting  piece,  c.r.,  cytoplasmic 
remnant,  g.a.,  anterior  granules.  g-P->  posterior  granules,  an., 
annulus.     m.p.,  main   piece,     e.p.,  end   piece. 


other  parts  of  the  spermatid  than  the  nucleus.  For  the  lit- 
tle we  can  do  in  the  way  of  giving  objectivity  to  this  general 
statement  we  will  first  examine  the  nearly  mature,  typical 


Evidence  from  Metazoan  Germ-Cells  5 

mammalian   spermatozoon    (fi^aire   36)    of    the   Fur   Seal. 

For  the  origin  of  the  various  parts  we  will  make  use  of 
the  summary  given  by  Ballowitz   (figure  37).      Concerning 


FIGURE    37.        "yOUXG"     SPERMATID     ( AFTER    BALLOWITZ). 

1,  nucleus.     2,  centrioles.     3,  idiosonie.     4,  mitochondrial  body 
chromatoid  body.     6,  spindle  remnant. 


5, 


the  nucleus,  i,  "it  is  certain  that  in  all  animals  the  chroma- 
tin-containing  part  of  the  sperm  proceeds  from  thu 
nucleus."  ^  On  this  point  there  is  agreement  among  observ- 
ers ;  so  whatever  may  be  the  truth  about  the  sperm-chromatin 


-rt 


FIGURE    38.       SPERMATID    OF    FUR    SEAL    ( AFTER    OLIVER ). 

n.,  nucleus,     c,  centrioles.     a.f.,  axial  filament. 


as  the  physical  basis  of  heredity  for  tlie  adult  animal,  there 
is  no  question  about  its  being  such  for  the  liead  of  tlie 
spermatozoon  itself. 


6 


The   Unity  of  the  Organism 


The  centrioles  {2,  figure  37)  are  "almost  always  double  and 
occupy  a  place  in  the  spermatid  close  beneath  the  surface  of  the 
cell."  ^  The  pair  is  typically  so  placed  that  the  axis  joining  them 
is   perpendicular   to   the   surface   of  the   spermatid^   the   member 


s- 


FIGURE    39.       SPERMATID   OF    FUR    SEAL     ( AFTER    OI.IVEr). 

h.c,   head  cap.     n.,   nucleus,     p.c,  proximate   centriole. 
centriole.     a.f.,  axial  filament,     s.,  remnant  of  sjihere. 


d.c,   distal 


toward  the  surface  being  known  as  the  distal  centriole  and  the 
one  toward  the  center  of  the  spermatid  the  proximal  centriole 
(figure  39  d.c.  and  p.c). 

As  to  the  part  played  by  each  of  these  in  the  development  of 


FIGURE    40.       SPEBMATID   OF    FUR    SEAL    (aFTER    OLIVER). 

n.,  nucleus,     c.h.,  chromatin  granules,     c.t.f.,  caudal  tube  filaments. 


the  sperm  we  learn  that  after  migrating  inward  until  they 
come  to  lie  very  near  the  nucleus,  if  not  in  actual  contact  with 
it,  the  centrioles  begin  their  development  with  the  following  out- 
come: "The  proximal  centriole  .  .  .  divides  into  two  portions, 
closely  adherent  to  the  nuclear  wall,  each  connected  by  a  fila- 
ment to  one  distal  group.      The  distal  centriole  divides  into  an 


Evidence  from  Metazoan  Gcrm-Cclh  7 

anterior  and  a  posterior  }3ortion.  The  posterior  portion  becomes 
the  anniilus  {an.,  figure  .S(j)  while  tlie  anterior  one  divides  again, 
forming  the  Xoduli  i)osteriores"  *  (g-P-,  figure  'Ui).  Throughout 
their  career  these  bodies  or  granules  are  highly  stainable  with 
certain   dye-stuffs. 

Out  of  the  idiosome  (S,  figure  ti7)  which  as  a  rule  is  a  body 
"fiir  sich" — in  itself  alone — develops  the  ])erforat()riuni.  or  lnad 
cap  of  the  adult  sperm  {li.c,  figure  3()).  It  is  agreed  that  the 
mitochondria  {Jf,  figure  37)  in  the  spermatids  of  many  animals, 
particularly  of  many  vertebrates  "furnish  the  material"  ^  for  the 
spiral  found  in  the  connecting  piece  {c.p.,  figure  36)  of  the 
sperm.  Although  no  spiral  is  present  in  the  Seal  sperm  it  may 
be  represented,  according  to  Oliver,  by  numerous  graTiulcs  sur- 
rounding the  axial  filament  in  the  connecting  piece,  l^ut  while 
the  connecting  piece  of  the  Seal  sperm  seems  not  to  be  tyi)ical 
as  regards  the  spiral,  it  presents  another  structure,  the  caudal 
tube,  or  "manchette"  of  some  authors,  in  a  form  which  is  s])ecially 
instructive  from  our  standpoint.  In  the  adult  sperm  this  struc- 
ture is  a  thin  sheath  enveloping  the  cytoplasmic  ])art  of  the  con- 
necting piece  and  lying  in  close  contact  with  the  persisting  cell 
membrane.  The  point  of  special  interest  about  it  is  that  its 
persistence  in  the  completed  sperm  of  the  Seal  a))pears  to  be 
exceptional,  for  it  is  known  to  disappear  entirely  in  the  course 
of  development  of  the  sperm  of  several  other  mammals.  It  is  a 
transitory  or  embryonic  organ  in  some  species  of  sperm,  but  a 
permanent  one  in  other  species,  just  as  gills,  for  exam|)le.  are 
transitory  organs  in  the  ontogeny  of  some  species,  as  a  frog,  but 
are  permanent  in  others,  as  fish. 

The  development  of  the  tube  in  the  Seal  sperm  is  es])rcially 
favorable  for  observation.  "It  may  be  reajclily  followed."  writes 
Oliver,  "from  its  first  appearance  up  to  its  final  incorporation 
in  the  connecting  piece  as  a  peripheral  layer,  or  sheath."  ^  Here 
then  is  a  structure  having  all  the  essential  marks  of  devel- 
opment due  to  heredity  and  likewise  one  the  "physical  basis"  of 
which  has  been  carefully  observed.  Mentioning  a  long  list  of 
investigators  who  believe  in  the  "derivation  of  the  caudal  tube 
by  a  process  of  cytoplasmic  differentiation  alone"  Miss  Oliver 
fells  us  that  her  study  of  the  develo])ment  of  the  fur  seal  sperm 
is  a  complete  confirmation  of  this  view.  As  to  the  very  begin- 
ning of  the  tube  we  read:  "Shortly  after  the  centrosomes  and 
their  tail  filament  have  reached  the  nuclear  membrane  there 
appears  in  the  cj^toplasm  surrounding  the  axial   thread   a  series 


8 


The   Unity  of  the  Organism 


of  delicate  filaments  attached  to  the  nuclear  membrane.  The 
proximal  ends  of  these  arise  in  a  circle  around  the  basal  end 
of  the  nucleus  with  the  centrosomes  as  a  center^  while  their 
distal  ends  project  freely  into  the  cytoplasm."  ®  (figure  40  c.t.f.) 
These  filaments  "are  at  first  very  short  and  thin^  but  they  in- 
crease in  length  and  thickness  rapidly.  By  the  progressive  dif- 
ferentiation of  the  cytoplasm  between  them  they  soon  fuse  into 
a  hyaline  tube,  surrounding  the  axial  thread  and  open  at  its  lower 
extremity."  (figure  41,  c.t.)  The  capital  point  is  that  we  have 
here   a   well-defined   structure   the   development   of  which   is   in- 


an 


FIGURE     41.        SPERMATID    OF    FUR    SEAL     ( AFTER     OLIVER). 

h.c,  head  cap.     c.t.,  caudal  tube,     an.,   annulus. 


dubitably  proved  to  depend  primarily  on  parts  of  the  cell  other 
than  the  chromatin.  Indeed  no  one,  apparently,  has  pretended 
that  the  chromatin  takes  a  part  in  its  production,  for  even  those 
investigators  who  have  not  believed  that  it  arises  from  the  cyto- 
plasm alone  have  held  that  it  originates  from  the  membrane  of 
the  nucleus. 

About  the  chromatoid  body  (5)  and  the  "spindle  remnant"  (6) 
(figure  37) J,  little  need  be  said  in  this  connection  as  they  seem  to 
be  inconstant  structures  the  significance  of  which  is  in  much 
doubt.  Finally,  mention  should  be  made  of  the  fact  that  a  con- 
siderable portion  of  the  cytoplasm  is  cast  off  entirely  in  the 
ontogeny  of  the  sperm  of  many  animals,  as  for  example  the  seal, 


Exndence  from  Metazoan  Germ-Cells  9 

the  body  (c.  v.,  figure  36)  being  all  that  is  left  of  tlie  substance 
at  the  late  stage  represented.  So  uuieh  by  way  of  ilhistratiou  of 
the  portions  of  the  s])ermatid,  or  germ  of  the  sjKrniato/oon.  in 
the  vertebrated  animals,  for  des])ite  the  great  variety  in  struct- 
ural details  presented  by  the  s))('rm  of  this  part  of  the  animal 
kingdom  I  think  all  will  agree  that  so  far  as  concerns  the  chief 
))oint  being  made  in  this  discussion,  what  we  have  ))r('sont<'d  is 
true  of  the  whole  ])hylum. 

(b)  Illustrated  by  the  Ontogeny  of  an  Insect  Sperm 

We  will  now  examine  the  ontogeny  of  a  very  different 
type  of  speiTn,  from  another  portion  of  the  animal  kingdom, 
the  insecta.  The  particular  species  chosen  is  the  fowl-tick 
{Argas  miniatus).  The  investigation  made  use  of  is  by 
Doctor  D.  B.  Casteel. 

The  series  of  figures  (42  a,  h,  c,  d,  e,  f,  g,)  will  help  to 
an  understanding  of  the  remarkable,  almost  unique  sperm  and 
spermogenesis  in  this  animal.  Figure  42rt  shows  the  nearly  ma- 
ture primary  spermatocyte.  Especially  to  be  noted  are  the 
mitochondria,  mi.,  scattered  uniformly  through  the  cytoplasm, 
and  the  striated  layer,  *.  /.,  on  the  outer  surface  of  the  cell. 
This  layer  is  sharply  demarked  from  the  underlying  cyto})lasui. 
The  striae,  disposed  perpendicular  to  the  surface  of  the  cell,  are 
excessively  fine,  and  when  looked  at  in  situ  end  on  "suggest  the 
appearance  of  a  faceted  compound  eye  or  of  honey-comb."  ® 
Concerning  the  genesis  of  this  layer  Casteel  says,  in  a  personal 
letter,  that  the  layer  begins  to  appear  at  the  surface  and  grad- 
ually increases  in  depth  until  the  completed  state  shown  in  figure 
42a  is  reached.  "The  striae,"  he  says,  "appear  to  be  forming 
from  the  undifferentiated  cytoplasm  sheath." 

All  the  figures  from  42  b  to  g  have  to  do  with  the  transforma- 
tion of  the  spermatid  (figure  42b)  into  tiie  spermatozoon,  h'rom 
figure  42b  one  sees  that  the  striated  layer  has  disap})eared  on  one 
side  of  the  cell  and  thinned  out  greatly  in  a  smaller  area  on  the 
opposite  side;  that  the  nucleus,  n,  has  moved  to  the  surface  of 
the  cell  in  the  middle  of  the  arr-a  of  disajipearance  of  the  striated 
layer;  that  the  large  plasmosomes,  pi.  42a  have  almost  entirely 
disappeared  from  the  nucleus;  that  the  vesicular  bodies  r.h.,  as- 
sembled  for   the   most   part   in   the   vicinity   of   the   nucleus,   are 


mi. 


r.'^j-M  n. 


-S.I.. 


A 


m.r.'- 


rn.r. 


FIGURE    42. 
FIGURE    42.       DEVEI.OPMEXT    OF    SPERM    OF    ARGAS    MIXIATUS     ( AFTER     CASTEEl). 

a.,  rupture  point  of  outer  tube,     c.p.,  cilia-like  processes,     fl.,  flagel- 
luin.      f.p.,   finger-form   jjrocess.      g.e.,   gelatinous   envelop,     i.e.,  in- 


10 


nx>, 


FIGURE    42. 

vagination  cavity,  i.t.,  inner  tube,  mi.,  niitochondria.  ni.r.,  mito- 
chondrial ring-,  n.,  nucleus,  o.d.,  oil  droj^ict.  o.t.,  outer  tul»c.  ]>!., 
plasmosome,  s.l.,  striated  layer,  v.b.,  vesicular  bodies.  1).]).,  l)egin- 
ning  of  inner  tube. 


11 


12  The  Unity  of  the  Organism 

in  processes  of  degeneration  and  disappearance;  and  that  the 
mitochondria,  mi,  look  as  though  they  had  collected  into  two 
well-defined  spherical  masses  m.r.,  on  the  side  of  the  cell  oppo- 
site the  nucleus  and  near  the  small  area  of  thinned-out  striated 
layer.  As  a  matter  of  fact  these  two  apparent  mitochondrial 
masses  are  the  opposite  sides  of  a  ring  seen  in  optical  section. 

Transformation  of  the  general  form  of  the  spermatid  now  be- 
gins by  the  indentation  of  the  side  opposite  the  nucleus,  this 
going  on  to  produce  first  the  quarter-moon  shape  shown  at  i.e., 
figure  12c.  By  the  still  further  growth  and  narrowing  the  edges 
of  the  cup  finally  come  together  to  produce  the  elongated  cavity 
shown  in  figure  42d,  o.t.  This  is  the  beginning  of  the  outer  tube 
which  becomes  long  and  relatively  narrow  as  development  con- 
tinues, (figure  42e_,  o.t.^.  At  an  early  period  in  the  growth  of 
this  tube  the  striated  layer  which  naturally  becomes  shut  into  the 
tube  breaks  up  over  most  of  the  circumference  of  the  tube  into 
what  resembles  a  dense  layer  of  long  cilia.  However,  since  the 
processes  are  not  motile  and  later  dissolve  and  produce  a  gela- 
tinous mass  within  the  tube,  their  resemblance  to  cilia  is  only 
superficial.  This  account  of  the  fate  of  the  striated  layer  applies, 
as  previously  intimated,  to  most  of  the  circumference  of  the  outer 
tube.  But  the  small  thin  area  of  the  layer  opposite  the  nucleus, 
shown  in  figures  42d  and  42c,  retains  the  cilia-like  processes. 
This  persistent  basal  patcli  (b.p.,  figure  42d)  is  the  starting 
point  for  an  important  part  of  the  future  spermatozoon,  the 
"inner  tube"  so  called  by  Doctor  Casteel  (i.t.,  figure  42e). 

While  these  profound  changes  are  going  on  in  the  portion 
of  the  spermatid  opposite  the  nucleus,  a  stout,  somewhat  finger- 
like  process,  (f.p.,  figure  42d)  is  formed  on  the  nuclear  side  of 
the  spermatid,  into  the  base  of  which  the  nucleus  migrates.  In 
the  meantime  the  vesicular  bodies  have  entirely  disappeared,  and 
the  mitochondria,  no  longer  disposed  in  the  ring  of  earlier  stages, 
have  assembled  into  an  irregular,  rather  diffuse  mass  toward  the 
basal  patch  {mi.,  figure  42d). 

At  the  time  when  the  outer  tube  has  reached  its  maximum 
length  and  is  somewhat  coiled,  the  inner  tube,  starting  at  the 
basal  patch  previously  described,  begins  to  grow  into  the  cavity 
of  the  outer  tube.  This  growth  continues  until  the  inner  tube 
is  approximately  as  long  as  the  outer  tube.  Figure  42e  presents 
an  advanced  but  not  completed  stage  of  growth  of  the  inner  tube 
i.t.  In  reality,  according  to  Doctor  Casteel's  interpretation,  the 
inner  tube  grows  at  the  expense  of  the  outer  tube,  for  when  the 


Evidence  from  Metazoan  Germ-Cells  13 

two  are  of  nearly  equal  length  the  entire  spermatozoon  is  only 
about  half  as  long  as  it  was  before  the  inner  tube  developed. 
All  of  the  mitochondria  of  the  cell  are  drawn  into  tlie  inner  tube 
as  it  grows,  and  finally  form  a  deeply  staining  mass  at  the  distal 
end  of  the  tube  {mi.,  figures  42e  and  ^ag). 

During  these  transformatory  operations  tlic  nucleus,  greatly 
reduced  in  size  proportionally  to  the  sjiermatozoon  as  a  whole, 
has  left  its  former  place  at  the  base  of  the  finger-like  j)r()eess, 
and  been  making  its  way  along  the  wall  of  the  outer  tube,  bur- 
rowing through  the  gelatinous  layer  on  the  outer  })art  of  this 
tube  {n,  figure  4<2e).  This  migration  continues  until,  when  the 
distal  end  of  the  inner  tube  reaches  nearly  the  end  of  the  outer 
tube,  the  nucleus  lies  in  the  wall  of  the  outer  tube  opposite  the 
end  of  the  inner  tube  {n,  figure  Igf.) 

The  final  act  of  transformation  takes  place  after  the  sperm 
has  left  the  male  tick  and  lies  in  a  spermatophore  sac  within 
the  genital  ducts  of  the  female.  This  act  begins  with  the  per- 
foration of  the  end  {a.  figure  42f)  of  the  outer  tube  by  the 
inner  tube.  Through  the  opening  thus  made  the  whole  inner  tube 
finally  passes,  really  by  a  slipping  back  of  the  outer  tube,  so 
that,  the  eversion  of  this  latter  being  completed,  the  two  tubes 
constitute  one  continuous  tube.  By  this  act  of  turning  inside 
out,  the  finger-like  process  in  which  the  nucleus  formerly  lay 
(figure  42d,  f.p.)  is  brought  into  close  proximity  again  with  the 
nucleus. 

Some  of  the  details  of  the  final  steps  in  the  transformation 
Doctor  Casteel  has  not  yet  been  able  to  make  out ;  but  these 
are  of  little  consequence  for  our  discussion.  Nor  has  the 
act  of  fertilization  been  observed.  Going  on  the  usual  cri- 
teria, the  end  of  the  sperm  containing  the  nucleus  would  be 
regarded  as  the  head.  But  surprisingly  enough,  in  moving, 
the  opposite  end,  the  end  containing  the  mitochondria,  goes 
foremost. 

This  sperm  and  its  development  are  so  unicpie  as  con- 
trasted with  those  occurring  in  most  animal  groups,  that 
one  might  be  almost  inclined  to  question  whether  there  may 
not  be  something  wrong  here — whether  the  case  may  not  be 
one  of  diseased  growth,  or  the  result  of  manipulative  mal- 


14  The  Unity  of  the  Organism 

formation,  or  something  else.  Any  such  suspicion  is,  how- 
ever, completely  done  away  with  by  the  fact  that  much  the 
same  type  of  spermogenesis  is  known  to  occur  in  other  ticks. 
Casteel  cites  particularly  the  observations  of  Katharine 
Samson  on  the  development  of  the  sperm  of  Ixodes  ricinus 
and  Ornithodes  mouhata  as  furnishing  cases  to  which  that 
of  Argas  is  "in  many  respects  parallel." 

After  all  that  has  been  said  in  the  previous  pages,  it  is 
almost  needless  to  point  out  the  significance  for  our  general 
contention  of  this  remarkable  case  of  spermogenesis.  Were 
we  living  in  that  comfortable  era  of  life-philosophy  wherein 
theologians  studied  nature  for  the  purpose  of  proving  that 
everything  in  it  was  made  to  meet  some  human  need,  we 
could  easily  recognize  this  case  as  one  designed  expressly 
to  assist  man  in  refuting  the  false  dogma  of  Chromatinic 
Omnipotence  in  heredity. 

It  is  hard  to  imagine  a  developmental  process  in  which 
denial  of  form-determining  power  to  non-chromatinic,  even 
non-nuclear  parts  of  the  cell  would  be  a  greater  folly  than 
would  be  the  denial  of  cytoplasmic  "form-determination" 
in  the  production  of  the  striated  layer  of  the  spermatid,  or 
of  the  growth  of  the  outer  tube,  or  of  the  inner  tube,  or 
of  the  turning  inside-out  of  the  outer  tube.  In  fact,  by  far 
the  greater  part  of  the  astonishing  transformations  here 
gone  through  are  cytoplasmic,  the  nuclear  changes  being 
relatively  slight. 

It  may  be  worth  while  to  remind  the  reader  again,  so 
boldly  does  the  real  truth  about  hereditary  substance  stand 
out  in  this  case,  that  the  cardinal  evil  in  the  chromatin 
dogma  is  that  it  implies  the  denial  of  great  masses  of  the 
most  direct  observational  evidence  we  have  as  to  what  the 
physical  bases  of  heredity  may  be,  and  so  tends  to  detract 
attention  from  them.  We  may  predict  that  the  important 
research  which  has  made  known  this  unique  case  of  organic 
genesis  will  pass  almost  unnoticed  by  the  geneticists  of  our 


Evidence  from  Metazoan  Germ-Cells  15 

day.  Were  their  attention  called  to  it  they  would  probably 
frankly  say  that  they  have  little  interest  in  f^cnesis  in  tliis 
sense.  That  the  sperm  here  described  is  not  peculiar  in 
every  respect  to  the  species  Argas  minmtus  is  certain  from 
the  meager  comparative  information  we  possess,  as  Casteel 
has  shown.  Nevertheless  not  merely  general  analogy,  but 
strong  indications  contained  in  even  the  little  comparative 
knowledge  w^e  have  in  this  particular  case,  warrant  the  sup- 
position that  in  some  respects  the  sperm  of  the  species  would 
be  peculiar  to  the  species,  to  say  nothing  of  the  genus, 
family  and  so  forth.  The  development  is,  consequently,  due 
to  heredity,  and  the  cytoplasm  is  "inheritance  material" 
as  ascertained  by  direct  observation. 

Evidence  from  the  Ovum 

We  now  turn  to  the  ovum  to  see  what  can  be  learned  con- 
cerning hereditary  substance  in  the  development  of  the 
ovum  iself.  Attention  should  be  called  at  the  outset  to  tlie 
important  difference  between  the  sperm  and  the  ovum  in 
the  kind  of  specialization  in  each.  The  sperm,  it  will  be 
noticed,  is  far  more  specialized  for  its  own  particular  life 
than  is  the  ovum,  this  "particular  life"  of  the  sperm  con- 
sisting in  its  great  power  of  locomotion.  As  a  consequence 
of  this  difference,  the  ovum  as  an  entity  has  no  such  sharp 
distinction  from  the  ovum  as  a  geiTninal  element  as  has  the 
spermatozoon.  This  difference  is  expressed  in  one  way 
by  the  assertion  that  the  fertilized  ovum  is  the  individual 
organism  in  the  one-celled  stage  of  its  life.  No  such  state- 
ment is  ever  heard  about  the  spermatozoon  for  the  obvious 
reason  that  the  sperm  does  not  transform  directly  into  the 
embryo  as  does  the  egg.  From  the  absence  of  so  distinctive 
a  character  of  the  ovum  as  such,  it  happens  that  tlie  hered- 
ity of  the  ovum  is  not  so  distinguishable  from  tlie  heredity 
of  the  organism  of  whose  life  it  is  a  stage,  as  is  the  case 


16  The   Unity  of  the  Organism 

with  the  sperm.  Nevertheless  we  are  bound  to  recognize 
that  the  ^gg  no  less  than  the  sperm  has  hereditary  attri- 
butes of  its  own,  and  that  other  substances  than  chromatin 
play  a  demonstrable  part  in  the  production  of  these.  In- 
deed the  main  discoveries  concerning  what  in  an  earlier 
chapter  was  called  the  promorphology  of  the  ^gg  are  of  this 
sort.  There  is  one  kind  of  promorphology  that  is  of  special 
importance  to  the  present  stage  of  this  discussion.  I  refer 
to  the  kind  known  sometimes  as  "germinal  localization"  and 
sometimes  as  "organ  forming  substances"  in  the  ovum.  The 
idea,  expressed  in  a  sentence,  is  that  in  the  eggs  of  some 
animals,  portions  of  the  ^gg  destined  to  give  rise  to  par- 
ticular parts  of  the  future  embryo  are  visibly  different  from 
other  portions  before  cell  division  begins,  in  some  cases  even 
before  maturation  and  fertilization  occur.  According  to 
our  understanding  of  heredity,  these  distinguishable  por- 
tions of  such  eggs  are  themselves  hereditary  attributes  not 
only  of  the  animal  species  to  which  the  eggs  belong,  but  of 
the  eggs,  no  less  than  are  distinctive  morphological  features 
of  the  adult  animals  or  of  any  developmental  stages.  The 
study  of  these  attributes  of  eggs  is  peculiarly  interesting 
since,  belonging  to  germ-cells  par  excellence,  if  we  can  get 
obseiTational  evidence  on  both  their  origin  and  destination, 
we  shall  have  direct  evidence  that  one  and  the  same  substance 
is  determined  on  the  one  hand  by  heredity,  and  on  the  other 
is  a  determiner  in  the  strict  genetic  sense  of  hereditary  at- 
tributes yet  to  be  developed. 

(«)  Eggs  of  Ascidians — Tlie  Facts 

Because  of  the  great  importance  of  the  observations  of 
E.  G.  Conklin  in  this  field,  and  of  his  general  views  con- 
cerning the  bearing  of  his  observations  on  heredity,  we  shall 
make  his  work  the  center  of  our  examination.  One  of  the 
most  important  of  Conklin's  investigations  was  on  the  eggs 


Evidence  from  Metazoan  Germ-Cells 


17 


of  several  species  of  Ascidians.  Ainonfr  the  great  merits  of 
this  investigation  arc  the  facts  that  the  normal  living  eggs 
were  studied  with  great  care,  and  that  the  comparative 
method  was  employed   to  a   considerahle  extent. 


ca. 


c  rt. 


FiGURK  4:^.     i)i:vp:r,op,^tKNT  OF  AX   AsciDi.xN   viAi    (afikk  coxki.ix). 

en.,   chorion,      c.p.,    clear    jirotoplasni.      c.r..   crescent  of   niesodcrMial 

suh.stance.     jLi'.N'.,  ^x'rniinal   Ncsiclc.     |).l>.,  polar-  l)o<l\ .  p. I.,  jx-riplicral 

layer    of    ])rotopla.sni.      t.e.,    test    cells.      \  .li.,    yellow  luMnisj)licrc    of 
egg.     yk.,  yolk. 


Three  species  of  simple  Ascidians,  Ciona  intestifuiUs,  Mol- 
gula  manhattensis  and  Siyela  partita  furnished  the  ^'^^i:,^ 
for  the  investigations. 

We  will  begin  the  examination  by  quoting  from  the  sum- 


18  The   Unity  of  the  Organism 

mar}^  of  results,  and  then  use  these  as  the  basis  of  a  closer 
scrutiny  of  the  conclusions.  Under  the  head  "Organization 
of  the  Egg"  we  find:  "14.  In  the  ovocyte  of  Cynthia  partita 
there  is  a  peripheral  layer  of  yellow  protoplasm,  {p.  I.)  a 
central  mass  of  gray  yolk,  (yk.)  and  a  large  clear  germinal 
vesicle  (g.  v.)  which  is  eccentric  toward  the  animal  pole 
(figure  43a).  These  same  parts  are  present  in  the  eggs 
of  other  ascidians,  but  are  difFerentlv  colored. 

"15.  When  the  wall  of  the  germinal  vesicle  dissolves  at 
the  beginning  of  maturation  divisions  a  large  amount  of 
clear  protoplasm,  containing  dissolved  oxychromatin,  is 
liberated  into  the  cell  body.  This  clear  protoplasm  is  ec- 
centric toward  the  animal  pole  and  is  distinct  from  the  yolk 
and  peripheral  layer. 

"16.  Immediately  after  the  entrance  of  the  spermatozoon 
the  yellow  and  clear  protoplasm  flow  rapidly  to  the  lower 
pole,  where  the  yellow  protoplasm  collects  around  the  point 
of  entrance ;  the  clear  protoplasm  lies  at  a  deeper  level.  The 
yellow  protoplasm  then  spreads  out  until  it  coA^ers  the  sur- 
face of  the  lower  hemisphere.  This  flowing  of  protoplasm 
to  the  point  of  entrance  of  the  sperm  is  comparable  with 
what  takes  place  in  many  animals,  though  much  more  ex- 
tensive and  rapid  here  than  elsewhere  (figure  48b).  ^ 

*'18.  The  sperm  nucleus  moves  from  the  point  of  en- 
trance toward  the  equator  in  a  path  which  is  apparently 
predetermined.  This  path  lies  in  the  plane  of  first  cleavage 
and  the  point,  just  below  the  equator,  at  which  the  sperm 
nucleus  stops  in  its  upward  movement,  becomes  the  posterior 
pole  of  the  embryo.  The  median  plane  and  the  posterior 
pole  are  probably  not  determined  by  the  path  of  the  sperma- 
tozoon but  by  the  structure  of  the  egg-  All  the  axes  of  the 
future  animal  are  now  clearly  established  antero-posterior, 
right-left,  dorso-ventral. 

19.  The  yellow  protoplasm  "collects  into  a  yellow  cres- 
cent with  its  middle  at  the  posterior  pole  and  its  horns  ex- 


Eiidence  from  Mefazoan  Germ-Cclh  10 

tending  about  half  way  around  the  egg  just  below  the 
equator  (figure  43c.  y.h.)  This  position  it  retains  tlirough- 
out  the  whole  development,  giving  rise  to  the  muscle  and 
mesenchyme  cells  (figure  43  d.  cr.) 

"20.  .  .  .  At  the  close  of  tlie  first  cleavage  (figure  43d) 
the  nuclei  and  clear  protoplasm  move  {c.p.)  into  the  upper 
hemisphere  and  thereafter,  throughout  development,  this 
hemisphere  contains  most  of  the  clear  j)rotoplasm  and  gives 
rise  to  the  ectoderm. 

"21.  .  .  .  When  the  clear  protoplasm  moves  into  the 
upper  hemisphere  the  yolk  is  largely  collected  in  the  lower 
hemisphere.     This  yolk-rich  area  gives  rise  to  the  endoderm. 

"24.  The  chief  factor  of  localization  is  protoplasmic 
flowing;  cell  division  is  a  factor  of  subordinate  value."  ^ 

From  the  numbers  given  in  this  quotation  it  will  be  recog- 
nized that  only  a  portion  of  the  summing  up  of  results  is 
here  presented;  and  it  will  be  understood  that  each  item  in 
the  summary  represents  a  lengthy,  detailed  description  in 
the  body  of  the  memoir.  Tliis  summary  we  may  again  sum- 
marize as  follows :  By  the  time  the  first  division  of  the  egg- 
cell  is  completed  the  portions  of  the  Qgg  whicli  are  to  give 
rise  to  three  great  groups  of  tissues  of  tlie  future  animal 
are  distinguishable  from  one  another  by  definitely  visible 
attributes.  The  clear  protoplasm  situated  at  the  upper 
pole  of  the  cell  will  give  rise  to  the  external  epithelium  or 
skin;  the  yolk-laden  protoplasm  in  the  lower  hemisphere  will 
produce  the  epithelial  lining  of  the  digestive  tract;  and 
from  the  yellow^  protoplasm  gathered  on  the  surface  at  the 
lower  pole  will  come  most  of  the  muscle  and  connective  tis- 
sue of  the  animal. 


(?;)  ConliirVs  Interpretation 

Passing  now^  to  a  consideration  of  these  facts  in  relation 
to  heredity  we  must  not  neglect  to  notice  tlie  two-fold  aspect 


20  The   Unity  of  the  Organism 

of  the  question,  namely  that  we  have  to  do  on  the  one  hand 
with  attributes  of  the  egg  itself  which  are  results  or  termini 
of  the  kind  that  gave  rise  to  the  conception  of  heredity ; 
and  on  the  other  hand  with  these  same  attributes  not  as 
results  but  as  causes  or  forerunners  of  later-appearing  at- 
tributes, also  by  the  same  criteria  due  to  heredity.  That  the 
egg  attributes  with  which  we  are  dealing  are  due  to  heredity 
is  obvious  from  the  fortunate  circumstance  that,  as  already 
indicated,  the  observations  under  review  were  comparative 
to  considerable  extent.  Thus  we  have  this  general  com- 
parison of  the  unripe  eggs  of  the  three  species :  "In  the  liv- 
ing eggs  of  Cynthia  this  peripheral  layer  is  clear  and  trans- 
parent and  contains  uniformly  but  sparsely  distributed 
yellow  pigment,  which  seems  to  be  associated  with  these  small 
refractive  spherules.  ...  In  Ciona  and  Molgula  also  these 
three  areas  are  distinguishable  in  the  living  ^gg  before  ma- 
turation, but  not  so  clearly  as  in  Cynthia.  In  Ciona  the 
peripheral  layer  is  nearly  transparent,  the  j^olk  is  a  brown- 
ish red.  ...  In  Molgula  both  the  peripheral  layer  and  the 
germinal  vesicle  are  transparent,  while  the  yolk  is  gray, 
with  a  faint  lilac  tinge."  ^  As  an  example  of  a  very  definite 
statement  of  specific  difference  between  the  eggs  we  read, 
"In  Ciona  the  same  type  of  protoplasmic  movement  occurs 
as  in  Cynthia,  but  with  certain  minor  differences."  ^ 

The  evidence  brought  forward  by  Conklin  of  species  differ- 
ences in  eggs  is  not  restricted  by  any  means  to  Ascidians. 
In  some  of  the  gastropod  mollusca,  for  example,  similar  re- 
sults were  reached  by  a  very  notable  research  on  the  quanti- 
tative relations  of  various  Qgg  organs  of  several  species  of 
the  genus  Crepidula.  To  be  explicit  in  a  single  case  only, 
the  relative  volume  of  macromeres  8A-3D  to  3a-3d  was 
found  to  be  12.1:1  in  Crepidula  plana  and  59.3:1  in  C. 
convex.  From  a  long  series  of  determinations  of  difference, 
Conklin  remarks :  "One  cannot  study  the  eggs  of  different 
animals  without  being  much  impressed  with  the   fact  that 


Evidence  from  Metazoan  Gcrm-Cclh  21 

the  distribution  of  yolk  to  the  four  macromeres  is  lii^lily 
characteristic  of  different  species  and  orders."  **  Vet,  as 
we  shall  have  occasion  to  j^oint  out  later,  it  seems  certain 
that  facts  of  this  kind  have  deeper  meanings  than  Conklin 
has  appreciated. 

After  all  that  has  been  said  in  previous  ])ages  ab(nit 
specific  differences,  the  point  aimed  at  here  will  be  obvious. 
The  "decidedly  thicker"  layer  of  peripheral  protoplasm  at 
the  lower  pole  in  Ciona  than  in  Styela,  and  the  brownish 
red  yolk  in  Ciona  as  contrasted  with  the  white  yolk  tinged 
with  lilac  in  Molgula,  are  indeed  "minor  differences."  But 
species  differences  in  adults  and  in  all  other  stages  are  minor 
as  a  rule.  The  difference  between  the  transparent-white 
color  of  the  full-grown  Ciona  intestinalis  and  the  pale,  green- 
ish yellow  of  the  adult  Molgula  manhattensis  is  also  minor 
as  contrasted  with  the  "same  type"  of  organization  of  the 
two  animals.  But  it  is  just  this  being  true  to  type  as  re- 
gards minor  differences  in  living  beings  that  has  given  rise 
to  the  conception  not  only  of  organic  species,  but  to  a  large 
extent,  of  heredity  as  well ;  and  how  can  any  one  recognize 
the  differential  color  of  the  grown-up  Ciona  as  compared 
with  the  grown-up  Molgula  as  due  to  heredity,  but  refuse 
to  recognize  the  differential  color  of  the  eggs  of  the  two 
animals  as  due  to  the  same  cause  .'^ 

We  are,  then,  bound  to  accept  the  attributes  of  the  eggs 
as  the  results  of  heredity,  and  so  inquire  about  the  "i)hysi- 
cal  basis"  or  "bearers"  of  these  attributes.  And  this  brings 
us  again  to  the  main  issue  of  this  section:  Have  we  or  liave 
we  not  observational  evidence  that  any  other  substance  or 
substances  than  chromatin  contribute  to  the  production  of 
the  egg-phenomena  under  consideration.? 

Conklin's  conclusions  and  mode  of  reasoning  as  to  \\\v 
bearings  of  his  observations  on  heredity  are  so  crucial  tor 
the  point  now  engaging  us  that  we  must  examine  tlu'in  in 
considerable  detail.     Returning  to  the  summary  of  results 


22  The   Unity  of  the  Organism 

upon  which  we  have  already  drawn  we  read :  "26.  The 
organization  of  the  ovocyte  is  not  the  initial  organization. 
The  yellow  protoplasm  (mesoplasm)  of  the  Cynthia  egg  is 
probably  derived,  at  least  in  part,  from  sphere  material 
(archoplasm)  which  arose  from  the  nucleus  at  the  last 
ovogenic  division.  The  yolk  (endoplasm)  is  formed  by  the 
activity  of  the  'yolk  matrix'  (Crampton)  which  also  is 
probably  sphere  material.  The  clear  protoplasm  (ecto- 
plasm) is  derived  from  the  g;erminal  vesicle  at  the  first 
maturation  division.  Thus  many  important  regions  of  the 
egg  come,  at  least  in  part,  from  the  nucleus,  and  a  method 
is  thereby  suggested  of  harmonizing  the  facts  of  cytoplasmic 
localization  with  the  nuclear  inheritance  theory."  ^^ 

Again,  we  find :  "This  truly  remarkable  condition  in  which 
considerable  portions  of  the  cytoplasm  are  traceable  to  the 
nucleus  is  of  the  utmost  theoretical  importance.  From  all 
sides  the  evidence  has  been  accumulating  that  the  chromo- 
somes are  the  seat  of  the  inheritance  material,  until  now  this 
theorj^  practically  amounts  to  a  demonstration.  On  the 
other  hand,  all  students  of  the  early  history  of  the  egg  have 
observed  that  the  earliest  visible  differentiations  occur  in 
the  cytoplasm,  and  that  the  position,  size  and  quality  of 
the  cleavage  cells  and  of  various  organ  bases  are  controlled 
by  the  cytoplasm.  However,  in  the  escape  of  large  quanti- 
ties of  nuclear  material  into  the  cell  body,  and  the  forma- 
tion there  of  specific  protoplasmic  substances  we  have  a 
possible  mechanism  for  the  nuclear  control  of  the  cytoplasm., 
and  when,  as  in  the  case  of  the  ascidians  and  fresh  water 
gastropods,  these  substances  are  definitely  localized  in  the 
egg,  and  can  be  traced  throughout  the  development  until 
they  enter  into  the  formation  of  the  particidar  portions  of 
the  embryo,  a  specific  mechanism  for  the  nuclear  control  of 
development  is  at  hand,  and  the  manner  of  harmonizing  the 
facts  of  cytoplasmic  organization  with  the  nuclear  inher- 
itance theory  is  clearly  indicated/'  ^^ 


Evidence  from  Metazoan  Germ-Cells  23 

(c)  Critical  Examination  of  ConkUuH  Interpretation 

One  is  stinick  at  the  outset  of  a  critical  exainiiiation  of 
these  sentences  by  the  fact  that  what  is  chiinied  is  that  ob- 
servations have  been  made  wliereby  a  nietliod  is  "sufrfrested 
of  harmonizing  tlie  facts  of  cytophismic  h)cahzati()n  with 
the  nuclear  inheritance  theory";  and  that  tiiis  siiggestcd 
harmonization  of  sucli  localization  with  the  "nuclear  in- 
heritance theory"  (not  "cliromatinic  inlicritancc  theory" 
be  it  noticed)  is  the  final  count  in  a  mass  of  evidence  wiiich 
"practically  amounts  to  a  demonstration"  that  tlie  "chro- 
mosomes are  the  seat  of  the  inheritance  material." 

In  other  words  objective  facts  which  are  only  suggestive 
of  a  conclusion  as  touching  the  inheritance  role  of  the  xvhoU 
nucleus  rise  to  the  demonstrational  level  as  touching  tlic 
same  role  of  very  small  parts  of  the  nucleus. 

The  method  by  which  this  particular  piece  of  h:)gical 
sleight-of-hand  is  performed  is  easy  to  see,  for  though  va- 
ried to  meet  the  exigencies  of  the  special  case,  it  folh)ws  the 
general  sclieme  of  elementalistic  interpretation  with  which 
we  have  become  famihar.  In  the  first  place,  the  fact  that 
the  observations  furnisli  very  little  if  anv  direct  evidence 
that  the  chromosomes  cause  the  cytoplasmic  flowing  and 
localization  is  made  innocuous  as  evidence  against  the  chro- 
mosome dogma  by  assuming  that  the  flowing  and  localization 
do  not  themselves  come  under  heredity.  That  such  an  as- 
sumption is  implied  in  the  argument  seems  certain  fiom 
the  fact  that  Conklin  did  not  consider  that  tlie  general 
theory  of  chromosomes  as  the  bearers  of  heredity  which  he 
had  espoused  made  it  incumbent  u})on  him  to  take  cognizance 
of  the  fact  that  he  himself  had  in  reality  testified  that 
portions  of  the  nucleus  other  than  chromosomes  are  the 
seat  of  inheritance  material.  We  have  here  another  and  a 
very  notable  case  of  shielding  a  pi-evalent  theory  l)v  defini- 
tion; that  is,  of  shutting  relevant   but    inimical    facts   away 


24  The   Unity  of  the  Organism 

from  it  by  definition.  That  Conklin  should  have  been  un- 
wittingly led  into  this  is  the  more  surprising  in  that  his 
own  studies,  particularly  those  on  comparative  morphology 
and  physiology  of  the  eggs  of  several  groups  of  animals, 
have  added  largely  to  the  proof  that  many  of  the  gross 
attributes  of  the  eggs  themselves  are  subject  to  heredity. 
Indeed  no  biologist  has  expressed  more  positively  than  he 
the  conception  that  the  egg  is  the  individual  animal  in  one 
of  its  stages  of  development  ".  .  .  from  its  earliest  to  its 
latest  stage  an  individual  is  one  and  the  same  organism ; 
the  Qgg  of  a  frog  is  a  frog  in  an  early  stage  of  development 
and  the  characteristics  of  the  adult  frog  develop  out  of  the 

egg^  but  are  not  transmitted  through  it  by  some  'bearer  of 
heredity.' "12 

One  wonders  if  Conklin  really  would  maintain  that  only 
the  attributes  of  an  animal  in  the  adult  stage  of  its  life  are 
subject  to  heredity;  or  even  that  the  attributes  of  any  of 
its  developmental  stages  are  not  so  subject,  were  he  con- 
fronted with  the  question  in  this  form.  Although  I  have 
been  unable  to  find  statements  in  his  writings  from  which 
one  may  positively  infer  what  his  answer  would  be,  yet 
several  passages  can  be  brought  together  which  can  not,  I 
believe,  be  harmonized  with  one  another,  but  reveal  real 
contradiction.  Thus  in  The  Mechanism  of  Heredity,  al- 
ready cited,  we  find :  "Differentiation,  and  hence  heredity, 
consists  in  the  main  in  the  appearance  of  unlike  substances 
in  protoplasm  and  their  localization  in  definite  regions  or 
cells.  Such  a  definition  is  as  applicable  to  the  latest  stages 
of  differentiation,  such  as  the  formation  of  muscle  fibers, 
as  it  is  to  the  earliest  differentiations  of  the  germ  cells, 
and  the  one  is  as  truly  a  case  of  inheritance  as  is  the  other. 
In  short,  different  substances  appear  at  an  earlier  or  later 
stage  in  the  development  of  all  animals,  and  these  substances 
A|C^  are  then  sorted  out  and  localized;  this  is  differentiation 
^^^Jz^Jieredity :  see  above].     Physiological  division  of  labor  in- 


Evidence  from  Metazoan  Germ-Cells  25 

volvcs  morphological  division  of  substance;  sorting  out  of 
functions  implies  sorting  out  of  tlie  material  substratum 
of  functions."  ^^ 

If  cytoplasmic  sorting  out  and  localization  as  well  in  tlie 
earliest  as  the  latest  stages  of  development  constitute  dif- 
ferentiation "and  hence  heredity,"  how,  one  must  ask,  can 
the  hypothesis  tliat  "the  chromosomes  are  the  seat  of  the 
inheritance  material"  ever  "practically  amount  to  a  demon- 
stration"? The  only  way,  so  far  as  1  can  see,  to  reconcile 
these  two  statements  is  to  say  that  in  so  far  as  the  expres- 
sion "seat  of  inheritance  material"  means  anything,  both 
chromosomes  and  cytoplasm  are  such  seats,  and  hence  that 
neither  is  the  seat  of  it. 

But  it  is  not  enough  to  point  out  that  there  is  contradic- 
tion here.  We  must  try  to  discover  just  how  so  careful  a 
reasoner  as  Conklin  should  fail  to  detect  it,  for  we  may 
feel  certain  that  the  failure  is  due  not  to  mere  oversight  or 
carelessness  but  to  some  defectiveness  in  standpoint  or  gen- 
eral mode  of  procedure.  Conklin  fully  realizes,  as  our  quo- 
tations show,  that  movements  of  the  cytoplasm  go  far  to- 
ward determining  the  attributes  of  the  eggs  of  Ascidians 
and  many  other  animals.  But  the  following  makes  the 
recognition  still  more  definite:  "Undoubtedly  the  most  im- 
portant of  all  the  localizing  factors  so  far  recognized  are 
cytoplasmic  movements."  ^^ 

Assuming  that  our  contention  is  valid  that  these  localiz- 
ing factors  of  the  cytoplasm  are  inheritance  factors  (and 
the  virtual  admission  of  this  by  Conklin  in  one  of  the  two 
statements  which  we  hold  to  be  contradictorv  should  be 
noted)  we  have  still  to  see  by  what  facts  and  reasoning  Conk- 
lin reaches  the  view  that  his  observations  support  the 
theory  that  "chromosomes  are  the  seat  of  inheritance  mate- 
rial." The  observations  In  this  case  which  support  the 
chromosome  theory  are  that  the  three  kinds  of  cyto})lasni 
of  the  Ggg:  the  yellow  protoplasm  (mesoj)lnsm),  the  sjjlurc 


26  The   Unity  of  the  Organism 

material  (archoplasm),  and  the  clear  protoplasm  (ecto- 
plasm) come,  at  least  in  part,  and  at  one  time  or  another, 
from  the  nucleus.  It  is  not  claimed  that  the  chromosomes 
or  even  chromatin  can  be  observed  to  produce  the  cytoplas- 
mic localization  or  any  of  the  other  distinctive  features  of 
the  Qgg,  as  for  example  the  different  colors  characteristic 
of  different  animal  species ;  or  the  different  kinds  of  proto- 
plasm in  the  same  species.  Just  what  can  be  observed  as 
to  the  relation  between  the  chromosomes  and  the  several 
kinds  of  material  which  pass  out  of  the  nucleus  into  the 
cytoplasm  is  not  much  dwelt  upon  by  Conklin  in  this  investi- 
gation. The  clear  protoplasm  (ectoplasm)  he  describes  and 
figures  very  definitely  as  being  the  major  clear  mass  of  the 
germinal  vesicle  set  free  in  the  cytoplasm  as  the  nuclear 
membrane  dissolves  at  the  beginning  of  maturation.  The 
chromosomes  are  said  to  be  distinguishable  at  this  time  as 
numerous  small  deeply  staining  bodies.  They  can  be  ob- 
served to  collect  together  in  the  "center  of  the  nuclear 
area"  and  certain  things  can  be  made  out  about  the  shapes, 
and  arrangement  relative  to  one  another  of  the  individual 
chromosomes ;  but  nothing  is  recorded  to  indicate  that  they 
take  any  part  in  the  movements  of  the  ectoplasm,  much  less 
in  the  production  of  it. 

Concerning  the  relation  of  the  other  two  kinds  of  cyto- 
plasm, the  yellow  or  mesoplasm  and  the  sphere  material  or 
archoplasm,  to  the  chromosomes,  Conklin's  description  is 
still  more  meager.  An  investigation  by  Crampton,  however, 
has  given  particular  attention  to  the  relation  of  the  sphere 
material,  or  as  he  calls  it  the  yolk-matrix,  to  the  chromatin. 
Crampton's  results  are  the  more  significant  for  this  discus- 
sion in  that  Conklin  was  undoubtedly  acquainted  with  them. 
As  the  result  of  several  searching  tests  of  the  chemical 
character  of  the  yolk-matrix,  Crampton  says :  "In  Molgula, 
certainly,  these  granular  masses  are  not  chromatin  in  the 
proper  sense."  ^^    Although  Crampton  believes  the  substance 


Evidence  from  Metazoan  Germ-Cells  27 

called  by  him  yolk-niatrix  and  considered  by  Conklln  to  be 
the  same  as  wliat  he  calls  sphere  material,  arises  from  the 
nucleus,  this  belief  rests  not  on  direct  evidence  of  the  pass- 
age of  the  substance  from  the  nucleus  into  the  cytoj)la.Mii, 
but  on  the  facts  that  when  the  substance  is  first  seen  in 
the  oocyte  it  is  a  small  mass  situated  in  the  cell-bod v  or 
cytoplasm  close  in  contact  with  the  nuclear  wall,  and  that  it 
reacts  to  stains  and  digestive  fluids  in  the  same  wav  that 
certain  granular  contents  of  the  nucleus  react.  But  Cramp- 
ton  is  very  explicit  in  pointing  out  that  this  substance  in  the 
nucleus  is  not  chromatin,  at  least  of  the  ordinary  kind. 
Upon  treating  the  cells  with  digestive  fluids  it  disappears 
from  both  inside  and  outside  the  nucleus,  the  true  chromatin 
being  then  left  in  clear  view  as  fine  granules  in  the  nuclear 
reticulum. 

So  far  as  concerns  the  endoplasm,  then,  even  thougli  it 
be  derived  from  s})here  material  and  this  in  turn  from  the  nu- 
cleus, the  most  trustworthy  evidence  we  possess  is  to  the 
effect  that  its  primal  nuclear  source  is  not  chromatin.  In 
view  of  such  facts  as  these,  made  known  by  Crampton  and 
others,  through  what  reasoning  would  Conklin,  to  whom  the 
facts  are  familiar,  still  hold  that  taken  all-in-all  they  su}>- 
port  the  chromatin  dogma  of  heredity.^  Readers  wliose 
minds  have  become  sensitized  to  the  general  type  of  reason- 
ing which  pervades  nearly  all  elementalistic  theorizing  and 
makes  it  to  some  extent  fallacious,  will  readily  anticipate 
about  how  the  argument  will  run  in  this  case.  But  it  will 
be  profitable  to  see  it  in  actuality.  After  saying  that  ^'sonie 
of  the  important  cytoplasmic  substances  can  be  actually 
seen  to  come  from  the  nucleus"  but  that  "this  does  not  indi- 
cate that  these  substances  exist  from  the  beginning  in  the 
nucleus;  on  the  contrary  there  is  direct  and  visible  evidence 
that  they  arise  epigenetically,"  Conklin  continues :  **Such 
epigenesis,  however,  does  not  signify  lack  of  primary  organ- 
ization;  on  the  other  hand  all  the  evidence  favors  the  vlrw 


28  The   Unity  of  the  Organism 

that  hack  of  the  organization  of  the  cytoplasm  is  the  or- 
ganization of  the  chromosomes,  which  is  definite,  determinate 
and  primary."  ^^      (Itahcs  mine). 

There  you  have  it  again !  Although  it  is  freely  granted 
that  you  can  see  the  cytoplasm  in  the  very  act  of  arising 
epigenetically  and  moving  about  to  become  definitely  located, 
that  is,  to  become  organized,  still  what  you  see  is  no  part  of 
the  real  essence  of  the  business.  "Back"  of  this,  in  the 
chromosomes,  which,  be  it  specially  noticed,  can  not  be  seen 
to  take  any  active  part  in  the  operations,  we  must  conceive 
is  the  "organization"  which  is  "definite,  determinate  and 
primary'' — in  other  words  which  is  The  Ultimate  Cause,  so 
far  as  heredity  is  concerned.  Again  I  repeat,  wearisome 
as  the  iteration  has  become,  that  the  fallacy  in  this  sort 
of  reasoning  is  not  in  holding  that  there  is  some  causal 
power  "back"  of  the  phenomena  to  be  explained,  but  that 
all  such  power  is  located  there.  Tliat  is,  stating  the  general 
point  in  its  application  to  the  special  case,  the  fallacy  lies 
not  in  holding  that  the  chromosomes  contribute  something 
to  the  hereditary  attributes  of  the  ascidians  and  other  ani- 
mal groups  whose  development  Conklin  investigated,  but  in 
the  implied  denial  that  the  cytoplasm  contributes  anything 
to  it. 

Conklin  probably  would  not  admit  that  there  is  real 
contradiction  between  the  observations  by  liimself  and  oth- 
ers, on  the  part  played  by  cytoplasm  in  the  early  stages  of 
development,  and  his  contention  that  the  evidence  now  "prac- 
tically amounts  to  a  demonstration"  of  the  correctness  of 
the  theory  that  the  "chromosomes  are  the  seat  of  the  in- 
heritance material."  What  he  would  probably  contend  is 
that  the  observations  are  opposed  merely  to  the  extreme 
form  of  the  chromosome  theory.  Thus,  speaking  from  an 
angle  of  the  general  subject  a  little  different  from  that  of 
cytoplasmic  localization,  he  writes :  "This  conclusion  is  not 
a  refutation  of  the  nuclear  inheritance  theory,  but  it  is  a 


Evidence  from  Metazoan  Germ-Cells  29 

profound  modification  of  it."  ^"^  And  still  more  recently 
he  has  said,  "Many  biolo^^ists  maintain  that  the  nucleus  and 
more  particularly  the  chromosomes  are  tlie  exclusive  seat  (jf 
the  'inheritance  material'  and  that  all  the  'determiners'  of 
adult  characters  are  located  in  them.  Against  the  extninc 
form  of  this  theory  many  general  and  s])ecific  objections 
may  be  urged.  General  objections  are  based  upon  the  con- 
sideration that  the  entire  cell,  cytoplasm  as  well  as  nu- 
cleus, is  concerned  in  differentiation  and  that  neitiier  is 
capable  of  embryonic  development  in  the  absence  of  the 
other.  Differentiation  is  indeed  the  result  of  the  interaction 
of  nucleus  and  cytoplasm,  and  how  then  can  it  be  said  that 
the  nucleus  is  the  only  seat  of  the  inheritance  material.''"  ^* 

An  elaborate  discussion  of  whether  the  language  here  used 
can  be  harmonized  with  the  statement  (juoted  above  about 
the  demonstration  of  the  correctness  of  the  chromosome 
theory,  by  saying  that  the  views  expressed  in  the  last  quota- 
tion merely  involve  a  "profound  modification  of  the  nuclear 
inheritance  theory"  would  smack  too  much  of  pure  dialectics 
to  deserve  a  place  in  this  volume.  Our  sole  concern  is  with 
the  truth  about  the  thing  itself. 

Conklin's  position  would  be  so  far  satisfactory  if  he 
would  permit/us  to  understand  his  statements  "the  entire 
cell,  cytoplasm  as  well  as  nucleus,  is  concerned  in  differentia- 
tion," and  the  one  about  modification  of  the  nuclear  inheri- 
ance  theory,  to  mean  that  cytoplasm  is  "inheritance  mate- 
rial" and  contains  "determiners."  Evidence  that  cytoplasm 
contains  "determiners"  is  even  more  jjositive  tlian  is  that 
for  the  theory  that  chromosomes  are  the  seat  of  such  things, 
for  the  simple  reason  that  we  can  observe  abundantly  cyto- 
plasm.in  the  act  of  producing  hereditary  structures,  wiiereas 
we  rarely  observe  chromosomes  operating  directly  in  this 
way.  But  such  permission  would  not,  I  fear,  be  forthcoming. 
If  it  would  be,  one  is  at  a  loss  to  understand  why  the  terms 
"hereditary  substance,"  "physical  basis   of  heredity,"   "de- 


30  The   Unity  of  the  Organism 

terminers,"  "factors"  and  the  like,  constantly  used  in  con- 
nection with  the  chromosomes  are  never  used  in  connection 
with  cytoplasm.  Indeed,  so  well  does  Conklin  present  the 
general  argument  for  the  participation  of  cytoplasm  in 
the  development  of  hereditary  structures  that  it  is  surpris- 
ing, not  to  sa}'-  disappointing,  to  find  him  neglect  to  present 
the  most  specific  argument  we  have  to  the  same  effect,  name- 
ly that  the  genesis  of  a  vast  range  of  such  structures  can 
be  directly  observed  to  be  largely  due  to  cytoplasmic  trans- 
formations. 

One  other  passage  in  Conklin's  general  argument  is  so 
significant  that  we  must  reproduce  it.  "Differentiation  is 
indeed  the  result  of  the  interaction  of  nucleus  and  cyto- 
plasm, and  how  then  can  it  be  said  that  the  nucleus  is  the 
only  seat  of  the  inheritance  material.'^  If  held  rigidly,  this 
theory  involves  the  assumption  that  the  cytoplasm  and  all 
other  parts  of  the  cell  are  the  products  of  the  chromosomes, 
and  that  therefore  the  chromosome  and  not  the  cell  is  the 
ultimate  independent  unit  of  structure  and  function ;  an 
assumption  which  is  contrary  to  fact.  Furthermore,  since 
heredity  includes  a  series  of  fundamental  \atal  processes 
such  as  assimilation,  growth,  division  and  differentiation, 
there  is  something  primitive  and  naive  in  the  view  that  this 
most  general  process  can  be  localized  in  one  specific  part  of 
the  cell,  something  which  recalls  the  long-past  doctrine  that 
the  life  was  located  in  the  heart  or  in  the  blood,  or  the 
ancient  attempts  to  find  the  seat  of  the  soul  in  the  pineal 
gland  or  in  the  ventricles  of  the  brain."  ^^ 

This  passage  contains  several  well-sent  shafts  not  only 
against  chromosomal  element alism,  but  against  the  elemen- 
talist  standpoint  generally.  And  I  must  recur  again  in  con- 
nection with  it,  while  the  facts  of  egg  organization  as  pre- 
sented by  Conklin  are  fresh  in  mind,  to  the  perception, 
indicated  in  previous  chapters,  that  the  physical-chemistry 
conception  of  the  cell  must  be  extended  to  the  organism.     If 


Evidence  from  Metazoan  Germ-Cells 


31 


Conklin  once  sees  the  full  force  of  tliis  contention,  he  will, 
we  may  hope,  be  ready  to  let  go  entirely  of  the  idea  that 
the  facts  of  cytoplasmic  organization  must  be  "harmonized 
with  the  nuclear  inheritance  theory."  ^^  He  will  then  see 
that  there  is  no  more  necessity  for  harmonizing  the  facts  of 
cytoplasmic  organization  witli  the  nuclear  inheritance  tlie- 
ory  than  there  is  for  harmonizing  the  facts  of  nuclear  or- 
ganization with  the  theory  of  cytoplasmic  inheritance. 


REFERENCE  INDEX 


1.  Ballowitz     255 

2.  Wilson,  E.   B.    ('00) 135 

3.  Ballowitz     277 

4.  Oliver     489 

5.  Oliver     479 

6.  Casteel     646 

7.  Conklin    ('05)    Ill 

8.  Conklin    ('05)    11 

9.  Conklin    ('05)    21 

10.  Conklin    ('05)    114 


11.  Conklin  ('05)  101 

12.  Conklin  ('08)  90 

13.  Conklin  ('08)  92 

14.  Conklin  ('05)  102 

15.  Crampton    ('99)    48 

16.  Conklin  ('05)  101 

17.  Conklin  ('08)  98 

18.  Conklin  ('15)  162 

19.  Conklin  ('15)  163 


Chapter  XV 

EVIDENCE  FROM  SOMATIC  HISTOGENESIS  IN 
MULTICELLULAR  ORGANISMS 

Ayl/ E  must  now  give  the  greatest  possible  concreteness 
^  '^  to  the  general  truths  stated  bj  Conklin  that  growth 
is  the  result  of  the  interaction  between  nucleus  and  cyto- 
plasm and  that  heredity  includes  such  fundamental  vital 
processes  as  assimilation,  grow^th,  division,  and  differentia- 
tion. But  the  one  and  only  way,  I  again  insist,  to  attain 
concreteness  and  certainty  in  the  matter  is  through  a  maxi- 
mum of  observation  coupled  with  a  minimum  of  inference. 
That  is,  the  goal  must  be  reached  mainly  by  direct  study  of 
the  anatomical,  histological  and  physiological  transforma- 
tions through  which  hereditary  attributes  are  produced.  The 
issue  can  be  met  squarely  only  by  a  still  further  considera- 
tion of  what  we  actually  know  about  the  participation  of 
all  sorts  of  elements  of  relatively  undifferentiated  cells  in 
the  production  of  obviously  hereditary  parts.  The  study 
of  interactions  between  nucleus  and  cytoplasm  and  of  as- 
similation growth,  etc.,  in  germ-cells  is  not  enough.  What 
we  have  seen  in  preceding  pages  about  the  development  of 
organs  in  the  protozoa  and  in  spermatozoa  is  that  much 
toward  the  end  sought.  Our  task  now  is  to  consider  the 
local  transformations  by  which  structures  are  produced  in 
multicellular  organisms,  especially  in  those  which  develop 
from  eggs. 

The  Mitochondrial  Theory  of  Heredity 

This  task  may  well  begin  by  examining  the  recent  efforts 
to  locate  the  "hereditary  units"  in  the  mitochondria  of  the 

32 


Evidence  from  Somatic  Histogenesis  33 

cytoplasm  instead  of  in  the  chromosomes.  This  effort  Is 
just  as  misdirected  as  is  the  effort  to  make  chromatin  the 
sole  hereditary  substance.  According  to  tlie  organismal 
conception,  all  life  phenomena,  including  those  of  inheri- 
tance, consist  in  the  activities  and  interactivities  of  an 
enoraious  number  of  substances  and  units  and  forces,  all  of 
which,  in  exhaustive  analysis,  arc  dependent  ui)on  the  organ- 
ism as  a  living  whole.  It  is,  therefore,  as  futile  to  hiuii  in 
one  comer  as  another  for  the  physical  basis  of  heredity 
in  an  exclusive  and  more  or  less  metaphysical  sense.  Any 
one  who  has  grasped  this  idea  will  know  beforehand  that 
the  proposal  to  make  mitochondria  fully  explain  heredity  is 
doomed  to  failure  no  less  certainly  than  was  the  proposal  to 
make  chromosomes  or  any  one  kind  of  cell  element  play  such 
a  role.  But  h3'potheses,  the  falsity  of  which  might  liave  been 
seen  before  they  were  tested,  may  still  be  useful.  If  those 
who  propose  them  can  be  convinced  of  their  fallacy  in  no 
other  way  than  by  testing  them  then  it  is  better  that  they 
should  be  tested  even  though  much  time  and  labor  be  given 
to  the  task.  Again,  the  evidence  brought  out  which  dis- 
proves an  hypothesis  may  be  highly  useful  in  establishing 
some  alternative  general  view.  This  result  has  been  espe- 
cially striking  in  the  case  of  the  mitochondrial  hypothesis  of 
hereditary  substance.  By  bringing  the  cell-body  back  into 
the  field  of  interest,  from  which  it  had  been  largely  excludi-d 
so  far  as  heredity  was  concerned,  by  the  nuclear  inheritance 
theory,  the  mitochondrial  hypothesis  has  resulted  not  merely 
in  proving  that  mitochondria  are  not  bearers  of  heredity 
in  the  elementalist  sense,  but  that  in  a  rational  sense  they 
are  sometimes  rather  closely  concerned  in  tlie  production 
of  hereditary  structures;  and,  of  even  greater  importance, 
that  still  other  cytoplasmic  material  is  likewise  so  concernrd. 
The  only  reason  why  the  mitochondrial  theory  of  hered- 
ity is  less  interesting  than  the  chromatin  theory  is  that 
there  is  so  much  less  obsei-vational  evidence  in  support  of  it. 


34  The  Unity  of  the  Organism 

In  fundamental  principle  the  one  is  no  more  and  no  less 
acceptable  than  the  other. 

Any  half  plausible  suggestion  that  a  particular  minute, 
obscure  part  of  the  germ-cell  may  be  a  "bearer  of  heredity" 
seems  to  endow  that  part  with  a  peculiar  fascination  to  biol- 
ogists who  haA'e  the  elementalist  habit  of  thinking,  and  this 
secures  to  it  an  inordinate  attention  until  the  untenability 
of  the  hypothesis  is  so  overwhelmingly  proved  that  even  the 
most  credulous  are  forced  to  abandon  it. 

Several  biologists  have  recognized  something  of  the  state 
of  mind  here  indicated  without,  however,  perceiving  its  real 
meaning.  Thus  in  a  review  of  the  work  of  Meves,  which 
will  be  examined  presently,  we  read:  "the  new  interpretation 
which  Meves  gives  at  this  time  indicates  that  many  are 
still  dissatisfied  with  the  all-sufficiency  of  the  [nuclear] 
theory,  and  are  eagerl}^  seeking  and  grasping,  as  it  were, 
the  first  visible  sign  of  any  other  substance  which  may  serve 
to  carry  the  hereditary  qualities."  ^  The  remark  to  be 
made  about  any  statement  of  this  kind  is  that  the  real 
though  usually  unperceived  ground  of  dissatisfaction  is  not 
with  the  all-sufficiency  of  the  nuclear  theory  of  heredity,  but 
with  the  all-sufficiency  of  any  theory  that  attempts  to  local- 
ize the  function  of  carrying  heredity  in  some  small,  specific 
fraction  of  the  germ-cells ;  and  that  the  attitude  which 
Doctor  Payne  well  characterizes  as  "eagerly  seeking  and 
grasping"  which  has  marked  so  much  of  recent  search  after 
the  physical  basis  of  heredity,  has  a  large  measure  of  genu- 
ine illusion  in  it.  Inspired  by  the  ages-old,  alluring  belief 
that  an  imperceptible  final  cause  and  explanation  is  hidden 
somewhere  within  or  behind  whatever  is  grossly  sensed,  the 
pursuit  becomes  "eager  and  grasping,"  which  is  another 
way  of  saying  that  it  becomes  more  or  less  irrational  and 
fitful. 

The  truth  of  these  remarks  is  rather  strikingly  exempli- 
fied in  the  short,  somewhat  feverish  history  of  the  mitochon- 


Evidence  from  Somatic  Histogenesis  35 

drial  theory  of  heredity.  The  meager  ()l)servati()iial  support 
for  the  hypothesis  that  tliese  ])artieuhir  cell  ineiiihers  are 
generally  bearers  of  heredity  relieves  us  from  the  necessity 
of  examining  it  in  any  such  detail  as  we  examined  the  chro- 
mosome theory.  But  there  are  several  things  al)out  it  of 
so  much  importance  that  we  must  look  into  it  somewhat. 
The  name  mitochondria  was  first  used  by  IJenda  for  a  "new 
cell  organ,  perhaps  serving  a  specific  function."  Benda's 
original  view  was  that  the  function  served  is  that  of  the 
motility  of  the  cell.  But  in  a  later  publication  he  presented 
observations  which  seemed  to  him  conclusive  proof  that  the 
mitochondria  of  the  sperm  are  situated  in  parts  of  it  which 
enter  the  Qgg  at  fertilization.  This  last  suggestion  gave 
an  added  impulse  to  the  study  of  the  bodies  occurring  In  the 
c3''toplasm  and  soon  many  new  names  were  applied  to  them, 
for  they  were  soon  found  to  present  differences  in  size, 
shape,  and  reaction  to  chemicals.  It  seems,  however,  that 
the  present  state  of  knowledge  justifies  us  in  applying  to 
them  all  the  one  term,  mitochondria,  though  without  Im]>ly- 
ing  that  they  are  all  exactly  alike.  They  may  be  held  to 
be  generically  alike  but  specifically  different. 

The  first  investigator  to  set  up  definitely  the  hypothesis 
that  cytoplasmic  elements  of  this  class  are  bearers  of  hered- 
itary qualities  seems  to  have  been  Meves. 

Only  a  few  of  the  very  many  investigations  since  devoted 
to  the  subject  can  be  noticed,  these  being  selected  for  their 
bearing  on  particular  aspects  of  the  problem.  In  the  first 
place,  the  position  of  ]\Ieves  himself  is  im])ortant.  Accej)t- 
ing  the  assum])tl()ns  formulated  by  ().  Ilertwig  in  1S7.")  of  a 
"substance  which  carries  over  the  begimiings  (Anlagen)  of 
the  parents  to  the  child,"  "  and  that  "this  substance  exists 
(in  the  germ-cells)  In  an  original,  histologically  uiidiffVren- 
tiated  condition";^  and  adding  his  own  reflection  that  not 
all  the  cytoplasmic  parts  of  the  spermatozoon  ( for  example, 
the  axial  fiber  of  the  tail)  possess  inheritance  potencies,  lie 


36  The  Unity  of  the  Organism 

advanced  the  hypothesis  that  the  mitochondria  answer  the 
conditions  of  Hertwig's  theory  for  the  cytoplasmic  part  of 
the  male,  though  probably  not  of  the  female  ge'rm-cells. 
Thus  Meves  reached  the  rather  attractive  conception  that 
^'heredity  is  accomplished  through  protoplasm  and  nucleus 
together."  ^  Were  we  to  know  no  more  than  this  about  his 
theory  we  might  suppose  his  position  to  be  that  of  a  genuine 
integrationist.  However,  his  very  next  sentence  does  not 
permit  us  to  question  any  further  the  orthodoxy  of  his  ele- 
mentalism.  "The  qualities  of  the  nucleus,"  he  says,  "are 
carried  over  by  the  chromosomes,  those  of  the  plasma  by 
the  chondriosomes."  ^  In  other  words  the  working  together 
of  nucleus  and  cytoplasm  which  he  conceives  is  not  of  the 
sort  which  makes  the  part  played  by  each  contribute  to  all 
the  results,  but  that  one  set  of  elements  produces  its  partic- 
ular part  of  the  total  effects,  while  another  set  produces 
another  part  of  the  effects. 

The  Mitochondrial  Theory  Tested  by  the  Ontogeny  of  the 

Spermatozoa 

The  utter  inadequacy  of  this  hypothesis  is  shown  by  some 
of  the  same  evidence  which  revealed  to  us  the  inadequacy  of 
the  chromatin  theory,  namely  that  when  we  come  to  study 
the  histogenetic  processes  by  which  innumerable  hereditary 
attributes  are  produced,  we  find  portions  of  the  cytoplasm 
other  than  the  mitochondria  taking  the  leading  part  in  the 
production.  Perhaps  no  single  one  of  the  many  instances 
examined  by  us  of  cytoplasmic  participation  in  the  produc- 
tion of  attributes  refutes  Meves'  hypothesis  more  complete- 
ly and  instructively  than  that  of  the  developing  sperm  of  the 
fowl  tick  already  described. 

We  have  already  examined  this  case  as  one  of  special  weight 
in  proving  that  the  cell-body,  in  contradistinction  to  the  nucleus, 
is   hereditary   substance.      Now  we   must  see  the  conclusiveness 


Evidence  from  Somatic  Histogenesis  37 

with  wliicli  this  particular  ontogeny  disposes  of  the  mitochondrial 
hypothesis  of  heredity  so  far  as  tiiis  case  is  concerned. 

Recurrence  to  the  descri))tion  and  figures  (42  a,  b,  c,  d,  e,  f )  will 
recall  that  the  mitochondria^  widely  distributed  through  the  cyto- 
plasm in  the  early  spermatocyte  {mi.)  assemble  into  a  rather 
sharply  defined  ring-shaped  mass  (m.r.,  figure  l-^b)  as  the  cell 
transformation  proceeds,  and  finally  take  a  position  in  the  devel- 
oping sperm  about  as  remote  as  they  can  get  from  some  of  the 
most  actively  and  fundamentally  changing  parts  of  the  organism. 
{mi.,  figures  42  e,  f,  g). 

There  is  no  more  possibility  of  explaining  the  development  of 
many  of  the  parts  of  this  sperm,  the  outer  sheath,  for  example 
{o.t.,  figures  36e  and  f )  as  due  to  the  influence  of  the  mitochondria 
than  as  due  to  the  influence  of  the  nucleus.  And  I  point  out 
again  for  the  hundredth  time  what  the  real  issue  is  here.  The 
observations  certainly  do  not  exclude  the  possibility  that  the 
mitochondria  exercise  some  invisible  influence  on  the  development 
of,  sav,  the  outer  sheath.  There  may  be  or  there  may  not  be  such 
an  influence.  But  the  observations  do  show  conclusively  that 
cytoplasmic  portions  of  the  cell  other  than  mitochondria  are 
operative  in  producing  the  outer  sheath. 

It  should  be  said  in  concluding  this  reference  to  tlie  sperm  of 
the  chicken  tick  that  Casteel  finds  the  mitochondria  located  finally 
in  the  end  of  the  sperm  opposite  that  which  contains  the  nucleus; 
and  that  this  end  goes  ahead  in  locomotion,  the  motion  being  pro- 
duced by  a  circlet  of  mobile  processes  at  this  end.  From  this  he 
believes  that  the  contractile  elements  are  mitochondrial  in  origin. 
The  conjecture  that  the  mitochondria  of  the  spermatocyte  take 
part  in  producing  the  motor  elements  of  the  sperm  tail  is  perhaps 
strengthened  by  the  observations  of  other  students,  notably  Lewis 
and  Robertson.  These  investigators  were  able  to  follow  tlie  mito- 
chondria in  the  ontogeny  of  the  living  sperm  directly  into  the 
tail,  where  they  transform  into  two  equal  threads  situated  along- 
side the  axial  filament. 

Tlie  Mitoclxondrial  Theory  Tested  hi/  Histogenesis 

If  this  hypothesis  that  mitochondria  in  developing  sperm 
cells  give  rise  to  the  motion-producing  structures  of  the 
sperm  tail,  then  the  mitochondria  would  be  genuine  "inlieri- 
tance  material"  for  these  particular  elements,  the  hereditari- 


38  The   Unity  of  the  Organism 

ness  of  the  motile  elements  being  especially  striking  in  the 
case  of  the  chicken  tick  sperm  from  the  fact  that  the  mode 
of  locomotion  in  this  spermatozoon  is  almost  unique.  But 
while  the  case  of  mitochondria  and  other  non-nuclear  parts 
of  the  cell  in  the  development  of  the  spermatozoa,  ought  to 
be  conclusive  that  although  mitochondria  can  not  be  "hered- 
itary units"  in  any  general  sense,  they,  as  well  as  other 
cytoplasmic  parts,  may  contribute  to  the  production  of 
hereditary  structures,  yet  it  would  not  be  so  accepted,  prob- 
ably, by  the  most  exacting  theorists  because  such  biologists 
would  not  allow  that  a  spermatozoon,  being  unicellular,  can 
have  organs  and  parts  which  are  subject  to  heredity  "in 
the  strict  sense"  (i.e.  in  the  sense  of  the  definition  of  hered- 
ity set  up  by  these  persons).  We  must,  consequently,  pro- 
ceed to  the  specific  task  of  this  section ;  namely  that  of 
considering  what  is  known  about  the  part  played  by  mito- 
chondria in  the  histogenesis  of  hereditary  structures  in 
multi-cellular  organisms. 

Nearly  all  the  studies  centered  upon  the  question  of 
whether  the  mitochondria  are  bearers  of  heredity  have  gone 
on  the  assumption,  quite  inadequate  according  to  my  view, 
that  the  problem  is  to  be  solved  by  ascertaining  whether  or 
not  the  bodies  are  persistent  cell  organs,  take  a  definite  part 
in  fertilization,  and  are  contributed  in  equal  quantity  by 
the  female  and  male  germ-cells.  In  other  words  the  assump- 
tion has  been  that  the  same  criteria  which  have  been  relied 
upon  to  prove  that  chromosomes  are  bearers  of  heredity, 
must  also  be  applied  for  deciding  whether  or  not  mitochon- 
dria have  the  same  office.  But  numerous  studies  have  also 
aimed  to  follow  the  mitocliondria  in  the  genesis  of  tissues, 
and  herein  lies  the  chief  importance  of  investigations  in 
this  domain.  Not  only  have  they  greatly  increased  our 
knowledge  about  the  role  played  by  various  parts  of  the 
cell  in  histogenesis,  notably  of  the  cytoplasmic  parts,  but 
they  have  put  us  in  possession  of  much  precise  information 


Evidence  from  Somatic  Histogenesis  39 

as  to  what  "hereditary  suhstancL'"  is. 

The  results  and  conclusions  tlius  far  reached  are  on  the 
whole  so  diverse,  often  so  conflicting,  that  any  attempt  at 
a  general  review  of  them  would  be  useless  for  this  discussion. 
However,  certain  of  tlie  results,  actual  or  claimed,  are  im- 
portant. For  example,  at  one  extreme  it  is  contended  that 
the  mitochondria  are  the  innnediate  })recursors  of  the  most 
distinctive  elements  of  all  classes  of  adult  tissues.  Thus 
Meves :  "All  these  differentiations  (of  embryonic  cells  into 
tissue  cells)  however  heterogeneous  they  may  be,  arise 
through  the  metamorphosis  of  one  and  the  same  elementary 
constituent  of  the  plasma,  the  chondriosomes.  The  chondri- 
osomes  are  the  material  substratum  lying  at  the  basis  of  the 
processes  of  differentiation,  which  become  the  specific  sub- 
stances of  the  different  tissues."  ^  As  an  extension  of  this 
view  we  learn  from  Lewis  and  Lewis  and  other  reviewers 
that  the  following  tissues  are  reported  on  the  authoritv  of 
a  long  list  of  workers  to  be  produced  by  the  mitochondria: 
fibrilla?,  myofibrilL'c,  fibrillje  of  epithelial  cells,  corneous  sub- 
stance, secretory  granules,  fat,  leuco-,  chloro-  and  chromo- 
plasts,  the  test  substance  in  foraminifera,  and  various  other 
tissue  elements. 

But  several  investigators,  notably  E.  V.  Cowdry,  have 
shown  the  inconclusiveness  of  the  evidence  on  which  the  con- 
tention is  based  that  neurofibrils  originate  from  mitochon- 
dria. "There  is  no  evidence,"  Cowdry  says,  "that  mito- 
chondria are  transformed  into  neurofibrils.  .  .  .  Tlie  mi- 
tochondria do  not  show,  either  by  a  variation  in  tluir 
morphology,  staining  reactions,  or  in  any  other  fashion, 
.  .  .  indications  of  being  transformed  into  material  of  dif- 
ferent chemical  composition."  *  Furthermore,  he  shows  that 
the  mitochondria  do  not  diminish  in  quantitv  in  anv  way 
commensurate  with  the  increase  of  neurofibrils,  as  thi'  ni'U- 
roblasts  transform  into  ganglionic  cells.  Eminently  worthy 
of   note,    as    bearing   on    our   contention    made   some    pages 


40  The   Unity  of  the  Organism 

back,  that  cytoplasm  itself  is  hereditary  substance,  is  Cow- 
dry's  detailed  description  of  neurofibrils  as  a  "differentia- 
tion of  the  ground  substance"  of  the  neuroblasts.  And  in 
a  later  paper  the  same  author  makes  a  strong  case  of  the 
view  that  while  mitochondria  are  "associated  in  some  way 
with  the  formation  of  many  substances,"  ^  it  is  highly  im- 
probable that  they  transform  into  them. 

On  the  whole  the  tendency  of  the  latest  investigations 
appears  to  be  to  deny  that  the  bodies  produce,  in  a  strict 
sense,  any  tissue  elements.  Thus  as  a  result  of  their  quite 
remarkable  studies  on  mitochondria  of  living  cells  Lewis  and 
Lewis  say,  following  the  enumeration  given  above:  "The 
above  theories  seem  impossible  to  correlate.  It  seems  evi- 
dent that  the  mitochondria  are  too  universal  in  all  kinds 
of  cells  to  have  the  function  of  forming  any  one  of  the 
above  structures  of  differentiated  tissue,  and  in  the  light  of 
what  cytological  chemistry  is  known,  it  appears  practically 
impossible  for  the  mitochondria  to  form  all  the  cell  struc- 
tures mentioned  above.  In  view  of  the  fact  that  the  mito- 
chondria are  found  not  only  in  almost  all  animal  cells  but 
in  plant  cells  as  well  it  seems  more  probable  that  they  play 
a  role  in  the  more  general  physiology  of  the  cell."  ^  The 
idea  that  the  mitochondria  are  primarily  concerned  with  the 
metabolism  of  the  cell  appears  to  be  gaining  ground  under 
the  present  comprehensive  and  critical  methods  of  investi- 
gation that  are  being  applied  to  them. 

The  Untenable  Hypothesis  that  the  Cytoplasm  of  the  Ovum 
is  Inheritance  Material  for  General  hut  not  for 

Special  Characters 

A  number  of  biologists  have  recently  put  forward  the 
hypothesis  that  while  the  cytoplasm  of  the  egg-cell  may  be 
"hereditary  material"  for  certain  of  the  general  attributes 
of    the    organisms,    chromosomes    "carry"    the    hereditary, 


Evidence  from  Somatic  Histogenesis  41 

of  tlio  more  specific  attributes.  This  conception  has  arisen 
from  a  considerable  range  of  observations  to  the  effect  that 
for  quite  a  time  in  the  early  ontogeny  of  many  animals  some 
of  the  attributes  of  the  embryo  can  be  seen  to  come  directly 
from  the  cytoplasm  of  the  egg.  Thus  both  Driesch  and 
Loeb  have  taken  special  notice  of  tlie  fact  tliat,  as  expressed 
by  Loeb,  "when  the  ]iroto])lasm  of  the  vgir  ])()ssesses  a  strik- 
ing pigment  the  larva  will  possess  tlie  same  for  some  time 
at  least";  and  that  "if  such  an  Qgg  is  hybridized  with  tlie 
sperm  of  a  form  whose  egg  is  unpigmented,  the  larva  will, 
of  course,  possess  a  'maternal'  quality  which  is  due  solely 
to  the  protoplasm  ( Driesch )".^^  And  in  the  same  connec- 
tion, Loeb  continues :  "It  is  obvious,  tlien,  that  during  the 
first  stages  of  development  an  influence  of  the  protoplasm 
upon  heredity  may  make  itself  felt,  which  will  disappear  as 
soon  as  the  protoplasm  of  the  egg  has  been  transformed  into 
the  tissues  of  the  embryo."  One  of  the  cardinal  questions  we 
have  to  consider  may  be  formulated  in  connection  with  tliis 
last  quotation:  Have  we  a  right  to  assume  that  because 
an  obvious  influence  of  the  protoplasm  upon  heredity  dis- 
appears on  the  transformation  of  the  protoplasm  into  tis- 
sue, therefore  all  such  influence  of  the  protoplasm  ceases.'^ 
To  answer  this  question  through  observations  upon  the 
protoplasm  of  the  cells  concerned  just  before,  during,  and 
just  after  the  transformations  is  exactly  the  central  aim 
of  this  section.  I  can  not  refrain  from  making  use  of 
another  sentence  from  Loeb  to  aid  In  defining  the  problem 
more  clearly.  "It  does  not  seem  to  me,"  he  writes,  "that  a 
discussion  as  to  the  relative  influence  of  protoj)lasm  and 
nucleus  upon  heredity  will  prove  very  fertile,  but  that  It  is 
necessary  to  transfer  this  problem  as  soon  as  possible  fi'oni 
the  field  of  histology  to  tliat  of  chemistry  or  physical  chcni- 
istry."  ^^  I  quite  agree  that  "discussion  as  to  the  relative 
influence  of  protoplasm  and  nucleus  ii])()n  heredity"  can  not 
be  very   fruitful.      But   the   grounds   of  my   ske])tlclsm   are 


42  The   Unity  of  the  Organism 

widely  different  from  those  of  Loeb.  According  to  my  view, 
the  question  is  not  one  to  be  settled  by  discussion  at  all, 
but  by  observation  coupled  with  a  measure  of  consistent 
reasoning.  Assuming  that  I  am  right,  to  "transfer"  the 
problem  "from  the  field  of  histology"  if  this  really  means, 
as  it  seems  to,  that  the  problem  should  be  taken  away  from 
histology,  no  matter  whether  to  the  field  of  chemistry  or 
any  other,  would  be  to  remove  it  all  the  further  from  ob- 
servation and  plunge  it  so  much  the  deeper  into  discussion. 
I  have  not  the  slightest  doubt  that  chemistry,  especially 
biochemistry  pursued  on  the  principles  of  physical  chem- 
istry, will  have  to  be  made  large  use  of  before  the  fullest 
possible  understanding  of  the  mechanism  of  heredity  is 
reached.  But  this  use  will  have  to  go  hand  in  hand  not 
only  with  morphological  studies  on  germ-cells,  but  as  well 
on  hosts  of  cells  during  the  whole  ontogeny.  Chemical  in- 
vestigation will  have  to  supplement,  it  cannot  supplant,  it 
cannot  even  lead,  histogenic  investigation.  If  there  is  one 
thing  made  more  positive  than  any  other  about  heredity  by 
modern  study  of  the  subject,  it  is  that  heredity  is  some- 
thing which  pertains  to  the  smaller  taxonomic  grades  of 
organisms,  races,  varieties,  species  and  so  forth.  It  would 
seem,  accordingly,  that  hardly  any  suggestion  for  the  study 
of  heredity  could  be  wider  of  the  mark  than  one  to  trans- 
fer it  from  the  only  field  which  makes  any  pretense  of  in- 
vestigating the  details  of  development,  and  taking  it  into 
a  field  like  that  of  physico-chemical  activity,  w^hich  is  no- 
toriously devoid  of  the  very  attributes  without  which  there 
would  be  no  such  thing  as  heredity.  Having  once  ascer- 
tained by  observation  as  much  as  possible  about  how  hered- 
itary attributes  are  actually  produced,  it  will  then  be  in 
order  to  leam  as  much  as  possible  about  the  chemistry  of 
the  processes.  Chemistry  can  do  its  share  in  solving  the 
problems  of  heredity  after  and  not  before  histogenesis  has 
done  its  share. 


Evidence  from  Somatic  Histogenesis  43 

Conklin  lias  expressed  more  definitely  than  any  other  biol- 
ogist with  whose  writings  1  am  acquainted,  the  idea  men- 
tioned above,  that  cytoplasm  "influences"  heredity  in  early 
ontogenetic  stages,  and  also  influences  adult  attributes  of 
the  major  taxonomic  groups,  but  })ecomes  inoperative  in 
the  later  stages  of  development,  the  heredity  of  these  being 
transferred  to  the  nucleus.  He  says,  "///  short,  the  c(j(j 
cytoplasm  fixes  the  general  type  of  development  and  the 
sperm  and  egg  nuclei  supply  only  the  details.^*  And  fur- 
ther: "We  are  vertebrates  because  our  mothers  were 
vertebrates  and  produced  eggs  of  the  vertebrate  pattern ; 
but  the  color  of  our  skin  and  hair  and  eyes,  our  sex,  stature, 
and  mental  peculiarities  were  determined  by  the  sperm  as 
well  as  by  the  ^gg  from  which  we  came.  There  is  evidence 
that  the  chromosomes  of  the  Qgg  and  sperm  are  the  seat  of 
the  differential  factors  or  determiners  for  ]\Iendelian  char- 
acters, while  the  general  polarity,  S3^mmetr3%  and  pattern 
of  the  embryo  are  determined  by  the  cyto])lasm  of  the 
iigg.''^  11  jf  ^^Q  points  in  this  last  quotation  be  viewed  in 
the  light  of  a  large  mass  of  relevant  evidence  not  usually 
taken  into  account  in  recent  discussions  on  lieredity,  and 
if  strict  consistency  in  the  use  of  terms  be  maintained,  the 
general  conclusion  will  be  quite  different  from  that  stated 
by  Conklin.  These  two  points,  the  concej)tion  of  "differen- 
tial" and  of  "determiner,"  must  now  receive  attention ;  but 
first  I  will  illustrate  my  position  by  a  case  presenting  the 
kind  of  evidence  to  which  I  have  referred. 

Species  Attributes  in  Single  Cells  of  Ail  id  t  Organisms 

In  general,  this  evidence  comes  from  the  field  of  histology, 
or  more  strictly  histogenesis.  The  most  convincing,  because 
the  most  direct,  evidence  from  this  source  is  that  })ertaiiiiiig 
to  the  development  of  hereditary  structures  in  adult  nieta- 
zoa    and    metaphyta.      The    structures    in    such    organisms 


44 


The  Unity  of  the  Organism 


which  are  the  most  indubitably  hereditary  are  those  which 
distinguish  the  smaller  but  yet  definite  taxonomic  groups. 
A  little  consideration  will  convince  one  that  about  the  most 
crucial  cases  would  be  those  in  which  the  development  of 
differential  attributes  could  be  traced  directly  to  cell  struc- 
ture and  development.  It  so  happens  that  vast  as  is  our 
knowledge  of  histogenesis,  the  part  of  it  which  answers 
directly  to  the  requirements  here  laid  down  is  by  no  means 
large. 

The  Spmules  of  the  Ascidian  Genus  Styela 

The  best  instances  I  have  been  able  to  find  are  super- 
ficial appendages  in  some  animals  and  plants,  so  small  that 
they  consist  of  a  few  cells  or  even  of  a  single  cell.  One  strik- 
ing instance  of  this  sort  has  come  to  light  in  my  own  studies. 


a. 


M.- 


-n 


FIGURE     44.        SPICULE     CELL    OF    STYELA    YAKUTATEXSIS     ( AFTER     HUXTSMAX). 

n.,  nucleus. 


It  is  that  of  the  minute  spines  which  cover  the  inner  surface 
of  the  branchial  siphon  in  some  species  of  the  ascidian  group 
of  Styalids.  Huntsman  first  called  attention  to  the  fact 
that  each  spinule  is  a  single  cell,  and  that  at  least  in  some 
cases  the  structures  furnish  differentiating  attributes  for 
species.  Miss  Forsyth  and  I  have  reexamined  the  point  for 
Styela  montereyensis  and  S.  yakutatensis,  and  are  able  to 


Evidence  from  Somatic  Histogenesis 


45 


confirm  Huntsman's  results.  As  Huntsman  luul  opportun- 
ity to  study  the  matter  in  a  larger  number  of  species  than 
Miss  Forsyth  and  I  have  had,  the  following  description  is 
taken  largely  from  his  paper.  Figures  44a,  1),  45,  46  are 
from  Huntsman  and  fimire  47  is  from  Hitter  and  Forsvth. 
By  comj)aring  figures  of  what  may  be  called  the  dorsal  view 
(figures  44a,  45,  46)  witli  the  side  view  (figure  441))  it  is 
seen  tliat  the  distinctive  feature  about  the  cell  whicii  con- 
stitutes the  spinule  is  a  shield-like  plate  on  one  side  of  the 
somewhat  elongated  cell,  the  distal  end  of  which  |)rojects 
more  or  less  beyond  the  cell  body,  the  whole  resembling  to 


FIGURE    45.       SPIXULE     CELT-    OF    STYELA    Pr.ICATA     (aKTKU     HUXl'S.MAX). 


some  extent  the  end  of  a  finger  with  its  nail.  The  shield 
is  harder  than  the  rest  of  the  cell,  and  is  probably  com- 
posed of  the  same  material  as  the  general  "test"  of  Ascid- 
ians,  animal  cellulose.  The  spinules  are  so  placed  that  the 
basal  portion  is  embedded  in  the  surface  layer  of  the  test 
on  the  inside  of  the  siphon,  the  shield  being  on  the  free  side 
of  the  cell  with  its  free  edge  ])ointed  toward  the  lumen  of 
the  siphon.  The  specific  attributes  furnished  by  the 
spinules  depend  upon  the  shape  and  structure  of  the  shields. 
The  free  edges  may  be  truncate  (figure  44,  S.  yakuta- 
tensis),  or  long-pointed  (figure  46,  S.  grech\i/i),  or  low- 
conical  (figures  45,  47,  ♦S'.  plicata,  S.  montercifcnsis). 
Again  the  edge  may  be  smooth  (figures  46,  47,  S.  grccUyi 


46 


The   Unity  of  the  Organism 


and  S.  montereyensis) ,  or  it  may  be  serrate  (figures  44,  45 
S.  yakutatensis,  S.  plicata). 

Viewing  this  case  in  the  hght  of  considerations  put  for- 
ward on  the  preceding  pages,  the  pertinent  queries  about 
the  heredity  of  the  shields  almost  ask  themselves :     What  is 


FIGURE    46.       SPIXULE     CELL    OF    STYELA    GREELEYI     ( AFTER     HUNTSMAN). 

the  "inheritance  material"  that  causes  the  sliield  to  be  short 
and  truncate  in  S.  yakiitatensis  and  long-pointed  in  S. 
greeleyi;  or  that  explains  the  serrated  edge  in  S.  yakiita- 
tensis and  S.  plicata  as  contrasted  with  the  smooth  edge  in 
S.  montereyensis  and  S.  greeleyi?  Is  the  "seat"  of  that  ma- 
terial in  the  cytoplasm  or  the  nucleus  of  the  shield-produc- 
ing cell?  Unfortunately  we  have  no  direct  obsei-i^ational 
infonnation    about    the    genesis    of   the    spinules.      But   the 


Evidence  from  Somatic  Histogenesis 


47 


indirect  evidence  whicli  bears  on  the  point  favors  over- 
wlielmingly  the  view  that  the  cytoplasm  is  chiefly  responsible 
for  the  shield.  Huntsman  supposes  the  sj)inules  to  be  de- 
rived from  the  cells  of  the  celhilose  tunic  characteristic  of 
ascidians.  He  may  be  right  in  this;  but  it  may  be,  too,  tljat 
they  are  derived  from  the  epithelial  lining  of  tlie  sipluni. 
The  matrix  of  the  cellulose  tunic  is  undoubtedly  largely  if 
not  wholly  produced  by  the  ectodermal  cells.  It  is  usually 
held   to   be   secreted   by   these   cells ;   but   in   some   cases,   in 


FIGURE    47.       SPIXULK     C'EIJ.    OF    S'l'YEI.A     MOXTEREYEXSTS     (aFTER    RFTTER     AND 

FORSYTH  ) . 

Perophora  for  example,  a  portion  of  the  cytoplasm  of  the 
cells  seems  to  become  transformed  into  the  cellulose.  The 
process  of  transformation  can  be  particularly  well  seen  in 
the  cells  which  line  the  branchial  siphon  of  developing  bhis- 
tozooids  as  shown  in  figure  32,  plate  III  of  my  memoir.' - 
The  ])arts  of  the  cell-bodies  turned  toward  the  cellulose  are 
here  long  drawn  out  and  the  protoplasm  gradually  becomes 
indistinguishable  from  the  surrounding  cellulose  substance 
in  which  it  is  imbedded.  If  now  we  imagine  this  proto})lasm 
to  transform  not  into  the  characteristic  cartilage-like  cellu- 
lose mass  spread  over  nearl}'^  the  whole  surface  of  the  body, 
that  from  innumerable  cells  fusing  into  a  connnon  mass,  hut 
each  cell  to  retain  its  individuality,  its  protoplasm  becoming 
the  shield  of  a  spinule,  we  should  have  what  these  styelids 


48  The  Unity  of  the  Organism 

actually  present. 

But  whatever  be  the  cells  which  transform  into  the  spin- 
ules,  all  the  available  evidence  indicates  that  the  cytoplasm 
is  the  chief  source  of  the  shield  part  of  the  spinule.  The 
possibility  is  not  excluded  that  the  nuclear  chromatin  also 
plays  some  part  in  the  development.  Indeed  investigations, 
notably  those  by  Duesberg  on  the  ascidian  egg  made  since 
"cytoplasmic  inclusions"  have  come  into  prominence  are 
distinctly  favorable  to  such  an  hypothesis.  But  this  is  very 
far  from  proving  that  such  chromatinic  influence,  assuming 
it  to  exist,  reduces  the  cytoplasm  to  pure  passivity. 

In  the  light  of  what  is  here  set  forth  let  us  examine  the 
view  expressed  by  Conklin  and  quoted  above  that  the  o^gg 
cytoplasm  fixes  the  general  type  of  development,  while  the 
nuclei  of  ^gg  and  sperm  together  "supply  only  the  details." 
The  examination  should  be  the  more  cogent  from  the  fact 
that  Conklin's  idea  was  based  largely  on  his  investigation 
of  ascidian  eggs,  some  of  which  pertained  to  the  very  same 
genus,  Styela,  as  do  the  spinules  just  described.  The  reader 
should  not  fail  to  notice  that  both  Conklin  and  myself  are 
dealing  with  single  cells,  he  having  to  do  with  egg-cells,  while 
I  am  concerned  with  spinule  cells.  Nor  should  the  fact  be 
lost  sight  of  that  these  two  categories  of  cells  stand  at 
about  opposite  extremes  in  the  life  career  of  an  individual 
Styela,  the  ^gg  being  at  the  very  beginning,  while  the  spinule 
is  produced  at  or  near  the  completion  of  adulthood.  Since, 
however,  both  cells  possess  attributes  distinctive  of  the 
species,  there  can  be  no  more  "filling  in  of  details"  at  one 
end  of  the  series  than  at  the  other,  so  far  as  concerns  the 
differential  attributes  under  consideration. 

Now  comes  the  main  point.  The  differential  attributes 
of  the  egg-cell  and  early  embryo,  their  "polarity,  symmetry, 
and  pattern,"  are  "determined  by  the  cytoplasm,"  to  use 
Conklin's  own  words.  What  is  the  evidence  that  these  attri- 
butes   are   thus    determined?      That   of   direct   observation. 


Evidence  from  Somatic  Histogenesis  49 

as  the  examination  of  Conklin's  work  has  shown.  But  if  on 
the  basis  of  such  observation  it  can  be  asserted  tliat  attri- 
butes of  egg  and  early  embryo  are  determined  by  the  cyt(j- 
plasm,  how  escape  asserting  that  the  same  sort  of  evidence 
touching  the  production  of  the  athilt  attributes,  those  per- 
taining to  spinule  cells,  are  likewise  determined  by  tlie  cyto- 
plasm ? 

It  was  remarked  above  tliat  if  all  known  relevant  facts 
were  taken  into  account  and  consistency  in  terminology  be 
maintained,  Conklin's  statement  to  tlie  effect  that  we  are 
vertebrates  because  our  mothers  were  vertebrates  and  pro- 
duced eggs  of  the  vertebrate  type,  but  that  our  species  and 
racial  characters,  color  of  skin  and  hair,  and  so  forth,  are 
determined  by  the  chromatin  of  both  parental  germ-cells, 
would  have  to  be  greatly  modified.  We  are  now  in  position 
to  see  what  modification  is  necessary.  Although  the  state- 
ment is  undoubtedly  true  that  we  are  vertebrates  because 
we  develop  from  vertebrate  eggs,  the  implication  that  the 
attributes  which  identify  us  with  the  human  species  and  the 
Caucasian  race  are  explained,  so  far  as  heredity  is  con- 
cerned, by  the  chromatin  of  the  geiTn-cells,  whether  male  or 
female  or  both,  is  not  in  accordance  with  all  the  observed 
facts  bearing  on  the  problem.  The  same  kind  of  evidence 
on  which  the  assertion  is  based  that  the  embryonic  charac- 
ters are  determined  by  the  cg^  cytoplasm,  requires  the 
assertion  that  skin  and  other  adult  characters  are  deter- 
mined by  the  same  means. 

This  leads  to  the  remarks  we  have  to  make  about  con- 
sistency in  the  use  of  terms.  The  critical  reader  will  hardly 
have  failed  to  notice  the  difference  in  application  of  flu- 
word  "determined"  as  used  by  Conklin  in  the  quotation 
we  are  examining.  When  it  is  said  that  the  color  of  skin, 
hair,  stature  and  so  on  are  determined  by  the  genn-cells, 
the  determining  act  or  condition  is  far  removed  from  that 
which  is   determined,   and   no   direct   causal   connection   be- 


50  The   Unity  of  the  Organism 

tween  the  two  is  established.  On  the  contrary  in  the  state- 
ment that  the  polarity,  symmetry  and  pattern  of  the  Qgg 
are  determined  by  the  cytoplasm,  the  determination  is  im- 
mediate and  observed.  Manifestly  in  a  literal  sense  "de- 
termined" is  proj^erl}'^  used  in  the  second  connection  but  not 
in  the  first.  The  cytoplasm  is  operative  on  the  spot,  so  to 
speak,  in  the  second  case.  It  is  concerned  with  an  immedi- 
ate result.  The  germ-cells  on  the  other  hand  are  not  really 
determiners.  They  are  not  concerned  with,  an  end  result, 
but  are  if  anything  instigators  of  a  long  developmental 
series  at  the  far  end  of  which  appear  the  attributes  in 
question:   skin,   color  and  the  rest. 

If  this  case  of  spinule  production  stood  alone  as  an 
instance  of  specific  characters  in  adult  animals  traceable 
to  cytoplasmic  activity  of  individual  cells,  it  would  be  a 
rather  small  base  on  which  to  erect  a  general  argument  in 
favor  of  cytoplasm  as  inheritance  material.  But  it  does 
not  stand  alone.  Indeed,  the  company  to  which  it  belongs 
will  almost  certainly  be  found  to  be  legion  when  systematic 
investigation  of  the  subject  shall  have  been  made. 

The  Spictdes   of  Sponges   and  Other  Invertebrates 

I  will  cite  a  few  more  cases.  In  several  widely  separated 
groups  of  animals,  spicules,  usually  either  calcareous  or 
silicious,  are  present  in  some  of  the  tissues.  These  are  often 
produced  by  one  or  a  very  few  cells,  and  often,  too,  their 
shape,  size,  and  probably  other  attributes  differ  from 
species  to  species  even  of  the  same  genus. 

The  spicular  system  reaches  its  greatest  development  and 
has  been  most  studied  in  sponges.  "The  spicules  of 
sponges,"  writes  Sedgwick,  "in  the  diversity,  symmetry,  and 
intricacy  of  their  form,  in  the  perfection  and  finish  of  their 
architecture,  constitute  some  of  the  most  astonishing  ob- 
jects in  natural  history."  ^^     Figure  48  gives  a  hint  of  the 


Evidence  from  Somatic  Histogenesis 


51 


facts  on  wliicli  this  statement  is  based.  Altliougli  the  pr()l>- 
lem  of  wliat  all  these  s])icules  are  for  does  not  dlreetly  con- 
cern us,  indirectly  it  does,  as  the  following  further  remarks 
of  Sedg^vick  will  indicate.  "While  it  is  pretty  clear,"  he 
says,  "that  the  main  function  of  the  skeletal  structures  is 
the  support  and  protection  of  the  sponge  body,  it  is  by  no 
means  easy  to  give  explanations  of  the  diversity  and  com- 


FIGURE    48.       SPICULES    OF    SPOXGES     (aFTEK    LAXKESTfUl) . 

plexity  of  form  which  they  present.  The  form  of  the  megas- 
cleres  is  probably  connected  with  the  form  of  the  canal  sys- 
tem with  which  they  are  in  relation  (F.  E.  Schulze)  ;  but  tlu" 
form  and  even  the  existence  of  the  microsclcres  defies  any 
reasonable  explanation."  And  then  comes  this  statement, 
highly  significant  for  almost  any  discussion  of  heredity : 
"By  some  spongologists  the  small  spicules  ari'  regarded  as 
functionless,  and  as  having  on  that  account  a  greater  value 
for  classificatory  purposes."  ^^ 

If  any  one  wishes  to  be  convinced  of  the  extent   to  whlilj 
the   spicule    forms    differ    with    different    species,    he   should 


52 


The   Unity  of  the  Organism 


consult  such  systematic  monographs  as  those  of  Schulze 
and  Sollas  in  the  reports  of  the  Challenger  Expedition.  A 
picture  of  such  a  group  of  spicules  as  that  shown  in  figure 
48  reminds  one  of  pictures  of  ice  crystals  he  has  seen;  and 
the  question  may  well  be  raised,  Are  not  these  spicules  in 
reality  crystallization  forms,  and  hence  as  devoid  of  hered- 
itary significance  as  are  snow  flakes?  The  fact  that  the 
form  they  have  depends  on  the  particular  group  of  sponges 
to  which  they  belong,  i.e.,  that  they  follow  the  rules  of 
biological  taxonomy,  is  very  strong  evidence  that  they  are 


FIGURE    49.       DEVELOPMENT    OF    A    SPICULE     (aFTER     LANKESTER). 

genuine  organic  productions,  and  not  mere  crystallizations. 
And  the  further  fact  that  they  follow  this  rule  even  though 
many  of  them  appear  to  have  no  functional  significance  in- 
dicates that  their  particular  forms  are  due  to  heredity  and 
not  to  modeling  by  extraneous  influences  in  each  individual 
sponge.  But  we  are  not  left  to  such  general  evidence  for 
support  of  the  supposition  that  they  are  true  organic 
products  and  subject  to  heredity.  Their  development  has 
been  studied  by  several  zoologists  and  the  results  leave  no 
doubt  about  their  nature  so  far  as  this  point  is  concerned. 

A  single  instance  will  be  enough  for  our  purpose,  but  it  should 
be  remarked  that  many  others  could  be  given.  This  is  taken  from 
the  excellent  summary  of  what  is  known  about  sponges  written 
by  Minchin  for  Lankester's  Treatise  on  Zoology.  "To  form  a 
triradiate  spicule  three  cells  migrate  into  the  parenchyma  from 


Evidence  from  Somatic  Histogenesis 


53 


the  dermal  epithelium  and  become  arranged  in  a  trefoil-like  figure 
(Figure  49).  The  nucleus  of  each  cell  then  divides  into  two,  in  sucli 
a  way  that  one  nucleus  is  placed  more  dee})ly  and  one  more  super- 
ficially. Between  each  pair  of  sister  nuclei  a  minute  sj)icule  ray 
appears_,  the  three  rays  being  at  first  distinct  from  each  other  but 


l-r.  Sys^. 


FIGURE    50 SEE    49. 

soon  becoming  united  at  the  center  of  the  system  (Figure  50  tr. 
syst.).  As  the  rays  grow  in  length  the  protoplasm  of  each  actino- 
blast  becomes  aggregated  around  each  of  the  two  contained  nuclei 
and  finally  more  or  less  completely  segmented  off  to  form  two 
formative  cells,  of  which  the  one  placed  more  internally  travels 
to  the  tip  of  the  spicule  ray^  while  the  other  remains  at  the  base 


FIGURE     51 SEE    49. 

b.f.c,  basal  formative  cell,     tr.syst.,  triradiate  system. 

(Figure  51,  b.f.c).  The  apical  formative  cell  sooner  or  later  dis- 
appears, returning,  apparently,  to  the  epithelium.  The  basal 
formative  cell  remains  at  the  base  of  the  ray  (Figure  51),  until 
this  portion  is  secreted  to  its  full  thickness.  It  then  migrates 
slowly  outwards  along  the  ray,  and  in  the  fully  formed  spicule  is 
found  adherent  to  the  extreme  tip." 


54  The  Unity  of  the  Organism 

That  such  a  mode  of  development  is  entirely  foreign  to  crystal 
production  hardly  needs  to  be  remarked.  But  if  further  proof 
to  the  same  effect  were  demanded^  one  other  strong  piece  of  evi- 
dence is  the  fact  that  most  of  the  spicules  are  not  composed  of 
inorganic  substance  alone  but  have  a  core  of  organic  matter. 

Although  the  technique  for  the  examination  of  cytoplasm  de- 
veloped during  these  last  years  has  not,  so  far  as  I  know,  been 
applied  to  the  spicule-producing  cells  of  sponges,  we  can  be 
reasonably  sure  from  the  study  of  other  secretory  cells  what  the 
general  results  will  be  when  such  application  sliall  have  been 
made.  They  will  bring  out  numerous  details  not  now  known  of 
how  both  the  nucleus  and  the  cytoplasm  act  during  spicule  pro- 
duction. 

Surely  it  is  not  necessary  for  me  to  dwell  again  on  the 
main  point  of  the  eyidence  here  presented.  The  nuclei  of 
the  spicule-fomiing  cells  may  take  an  active  part  in  pro- 
ducing the  spicules.  Indeed  from  our  general  knowledge  of 
nuclear  activities,  illustrations  of  which  were  given  in  an 
earlier  chapter,  it  is  probable  that  such  will  some  day  be 
demonstrated  to  be  the  case.  But  the  proof  of  nuclear 
activity  in  spicule  production  will  not  he  disproof  of  the 
already  observed  cytoplasmic  activity  in  spicule  production. 

Other  animals  that  may  be  mentioned  in  which  spicules 
are  produced  in  much  the  same  way  and  have  the  same  taxo- 
nomic  diversity  and  constancy  are  the  alcyonaria  among 
coelenterates,  the  holothurians  among  ecliinoderms,  and 
some  of  the  compound  ascidians,  particularly  of  the  family 
didemnids.  Relative  to  the  specificity  of  the  structures  in 
holothurians,  we  have  this  piece  of  significant  information: 
"These  calcareous  bodies  are  of  great  value  to  the  system- 
atist  in  classifying  the  smaller  groups,  such  as  genera  and 
species.  Although  their  general  characteristics  are  fairly 
similar  within  the  several  families,  the  different  shapes  of 
spicules  are  not  sufficiently  constant  to  be  used  as  diagnos- 
tic characters  of  such  large  divisions."  ^^  In  other  words, 
so  far  as  these  animals  are  concerned,  should  it  be  found. 


Evidence  from  Somatic  Histogenesis 


55 


as  it  almost  certainh'  will  be,  that  Hic  cjto])lasni  is  "hered- 
itary substance"  for  the  production  of  s])iciih's,  tlic  reverse 
of  Conklln's  generalization  that  the  cytophism  dctcrniiues 
the  larger  taxonomic  features  of  animals  while  chromatin  is 
the  seat  of  the  inheritance  factors  for  "fillinir  in  details" 
turns  out  to  be  true.  Species-marking  details  arc  just 
what  we  are  able  to  see  the  cytoplasm  fill  in. 

The  "Hairs''  of  Higher  Plants 

For  a  few  more  instances  of  species  characters  in  multi- 
cellular organisms  brought  down  to  single  cells,  we  turn  to 
the  plant  world. 


FIGURE    5-2.  FIGURE    53.  FIGURE    54-. 

FIGURE    52.       TRICHOIMES    OF   PAPAVER    ORIENTAI.E     (aFTER    CANNON ). 
FIGURE    53,       TRICIIOMES    OF    P.    PILOSUM     ( AFTER    CANNON  ). 
FIGURE    54.       TRICIIOMES    OF    P.    SO^IXIFEHr^I     (.MTrH    CAN  SOS  ). 

The  "hairs"  or  trichomes  borne  on  the  leaves,  flowers 
and  smaller  stems  of  innumerable  flowering  plants  are 
usually  composed  of  only  a  few  cells,  so  that  the  characters 
they  have  are  often  referable  to  the  individual  cells.  Can- 
non has  lately  investigated  these  structures  in  several  gr()U])s 
of  plants,  and  while  he  was  not  aiming  at  the  particular 
question  now  occupying  us,  some  of  his  results  are  (jiiife  to 
the  point  for  this  discussion.  An  exami)le  of  particularly 
distinct  specificity  of  the  hairs  Is  ])resented  in  the  following: 
"The  trichomes  of  the  three  species   |  of  p<>pi)y  |   are  similar 


56  The  Unity  of  the  Organism 

in  form  and  size,  but  they  are  unlike  in  quality  of  rough- 
ness. In  Papaver  orientate  (52  a,  b)  and  Papaver  pilosum 
(53  a,  b,)  the  distal  ends  of  the  superficial  cells  project  be- 
yond the  general  surface  of  the  trichome  and  turn  out  at 
a  rather  acute  angle.  In  Papaver  soTnniferum,  however, 
these  cells  did  not  extend  beyond  the  general  surface,  with 
the  effect  that  the  trichomes  of  this  species  are  smooth  (fig- 
ure 54  a,  b)."  ^^ 

This  is  especially  instructive  because  the  attribute  in 
question  pertains  to  the  shape  and  position  of  cells,  and  not 
to  differentiation  within  the  cells.  There,  consequently,  is 
no  room  for  even  a  reasonable  surmise  that  the  attribute  is 
explained  by  the  chromatin  instead  of  the  cytoplasm.  Even 
though  the  account  gives  no  information  about  the  position 
and  behavior  of  the  nuclei  of  the  cells,  it  is  hardly  con- 
ceivable that  any  one  would  maintain  that  the  substance 
itself  of  the  cell-tips  is  not  the  main  factor  in  the  out- 
turning  of  these  tips  characteristic  of  Papaver  orientate 
(figure  52  a,  b). 

Dealing  with  quite  another  type  of  trichomes,  those  of 
the  walnut.  Cannon  writes:  "A  character  which  easily  dis- 
tinguishes the  short  secreting  trichomes  of  Jwgtans  cati- 
fornica  from  those  of  Jugtans  regia  or  Jugtans  nigra 
is  the  length  of  the  head-cells."  ^'^  Both  drawings  and  tables 
of  measurements  of  the  heads  showing  lengths  and  diameters 
are  given  to  bring  out  the  positiveness  of  the  distinction. 
Furthermore,  details  of  cell  division  and  cell  structure  dur- 
ing the  development  of  the  trichomes  are  furnished ;  so  the 
visibte  evidence  that  the  cytoplasm  is  "inheritance  material" 
in  this  case  is  beyond  question.  What  the  invisible  evidence 
may  be  remains  for  further  investigation  to  discover,  but  of 
one  thing  we  may  be  sure:  no  matter  how  many  facts  of 
development  now  invisible  may  later  become  visible  they  will 
not  destroy  the  validity  of  the  present  visible  evidence. 

How  far  it  would  be  possible  to  go  on  pointing  out  specific 


Evidence  from  Somatic  Histogenesis  57 

characters  of  plants  that  are  referable  to  Individual  cells  I 
do  not  know;  but  judging  from  the  instances  that  come  to 
view  even  in  my  limited  knowledge  of  the  subject  It  might 
be  carried  to  an  almost  indefinite  extent.  Anotlier  instance 
which  I  recall  from  the  experience  of  my  student  days  is 
the  case  of  mosses.  The  serration  of  the  leaves,  I  remem- 
ber, was  one  of  the  features  relied  upon  for  generic  and 
specific  characteristics,  and  I  also  remember  that  the  indi- 
vidual teeth  often  if  not  always  consisted  of  one  or  a  very 
few  cells. 

Cell-wall  Structures  in  Higher  Plants 

That  the  cell-wall  is  a  structure  of  great  importance  in 
plants  is  known  to  everybody ;  and  the  veriest  tyro  in  plant 
histology  has  learned  something  of  the  enormous  variety 
and  definiteness  of  character  in  different  tissues  and  dif- 
ferent kinds  of  plants  presented  by  this  part  of  the  cell. 

A  very  brief  reference  to  two  plant  structures,  pollen- 
grains  and  wood  tissue,  will  be,  perhaps,  a  sufficient  re- 
minder of  what  there  is  for  us  in  this  domain.  A  typically 
formed  pollen-grain  is  a  minute  spheroidal  body  containing 
two  cells,  one  known  as  the  antheridial  or  germinative  cell 
and  the  other  as  the  sterile  or  vegetative  cell.  The  wall  of 
the  grain  consists  of  an  outer  coat,  the  exine,  and  an  inner, 
the  intine.  The  elaborateness  of  structure  which  these  coats 
may  reach  is  astonishing  if  regarded  in  the  crepuscular  liglit 
of  the  theory  that  cells  are  "simple"  things.  The  most  dis- 
tinctive thing  about  the  pollen-grain  is  the  pollen  tube 
which  is  produced  on  one  side  of  the  grain  and  througli 
which  the  antheridial  cell  reaches  the  ovule  in  fertilization. 
The  taxonomic  variety  which  is  our  main  interest  just  lure 
pertains  largely  to  the  sculpturing  of  the  surface  of  tlu' 
exine  and  to  the  structure  of  tlic  exine  at  the  point  where 
the  pollen  tube  will  break  through.  It  is  well  known  to 
botanists   that  the  "spikes,  warts,   ridges,   combs,  etc."   of 


58  The  Unity  of  the  Organism 

the  surface  of  the  grains  are  in  general  definitive  of  taxo- 
nomic  groups  of  plants.  And  concerning  the  places  of 
emergence  of  the  pollen  tube  we  read :  "The  number  of 
these  peculiarly  organized  points  of  exit  is  a  fixed  one  in 
each  species,  and  often  in  whole  genera  and  families."  ^^ 
As  to 'the  way  these  various  structures  are  produced  we  have 
this  very  definite  statement:  "The  sculpturing  upon  the 
outer  surfaces  of  the  spores  of  mosses  and  ferns  and  the 
corresponding  pollen  grains  of  the  Phanerogams  can  in 
most  cases  be  attributed  to  the  activity  of  the  protoplasm 
surrounding  the  developing  spores."  ^^ 

That  the  main  tissues  of  plants  present  taxohomic  char- 
acters is  amply  illustrated  in  the  wood-tissues.  The  facts 
concerning  these  tissues  have  been  almost  forced  into 
prominence  by  the  needs  of  fossil  botany,  though  they  have 
also  been  much  studied  as  a  part  of  ordinary  plant  mor- 
phology. The  section  on  fossil  wood  in  Zittel's  Handbook  of 
Palaeontology  is  a  considerable  resume  of  knowledge  in  this 
field,  and  contains  numerous  statements  and  figures  which 
bring  out  impressively  the  general  truth  of  the  specificity 
of  the  tissues  of  trees.  After  necessary  allowance  is  made 
for  the  strictly  botanical  unsatisfactoriness  of  many  of  the 
species  and  genera  recognized  by  palaeobotanists,  it  is  not 
doubted,  so  far  as  I  know,  that  on  the  whole  the  kinds  of 
tissue  they  describe  do  in  reality  represent  different  kinds 
of  trees.  A  single  illustrative  quotation  will  serve  to  make 
concrete  what  is  here  dealt  with  in  general  terms :  "The 
phloem  segments,  like  those  of  the  xylem,  are  divided  by 
few-seriate  pith  rays  into  rather  regular  two-  to  four- 
seriate  rows  of  cells  made  up  of  thin-walled,  small-celled 
elements  in  the  trunk  of  Zamia  floridana,  etc.,  and  Stan- 
geria.  But  ...  in  the  trunks  of  Cycas,  Dion,  Encepha- 
lartos,  Macrozamia  and  doubtless  most  cycads,  as  likewise 
in  the  Cycadeoideae,  sclerenchymatous  elements  are  more 
or  less   numerously   and   regularly   interspersed   among  the 


Evidence  from  Somatic  Histogenesis  59 

row  cells,  thus  giving  much  more  strength  to  the  stem,"  ^^ 
And  the  author  goes  into  considerable  detail  in  discussing 
the  presence  and  tlic  absence  of  "wood  tracheids,"  "scahiri- 
form  pittings,"  "border  pits,"  "spirally  tliickened  walls" 
and  "sieve  tubes"  in  tlie  genera  Zamia,  Cordaites,  Stanfjeria 
and  Cycas. 

That  this  well-nigh  endless  variety  of  character  of  the  cell- 
wall  in  adult  plant  tissues  is  due  to  the  activity  of  the  cell 
protoplasm  appears  never  to  be  questioned  by  botanists 
so  long  as  they  are  dealing  with  the  actual  structure  and 
development  of  the  wall.  "The  cell-membrane  is  produced 
by  the  protoplasm,"  we  read  in  the  section  on  morphology 
in  the  Lehrbtwh  der  Botanik  by  Strasburger,  Jost,  Schenck 
and  Karsten  (11th  German  edition)  this  section  being  from 
the  pen  of  Strasburger  himself.  Tliis  simple,  unqualified 
statement  of  fact  by  Strasburger  is  the  more  noteworthy 
because,  as  we  saw  in  another  connection,  he  has  been  one 
of  the  extremists  on  the  chromosome  dogma  of  heredity. 
Reading  his  statement  that  the  cell-membrane  is  produced 
by  the  protoplasm,  with  the  indubitable  fact  in  mind  that 
this  membrane  presents  innumerable  characters  which  are 
taxonomically  definitive,  and  hence  are  hereditar}^  according 
to  the  best  criteria  we  have  of  hereditary  characters,  it 
seems  impossible  to  avoid  seeing  that  it  implies  an  irrec- 
oncilable contradiction  of  Strasburger's  often-repeated 
view  that  the  chromosomes  are  the  sole  bearers  of  heredity. 

To  round  out  the  primarily  factual  part  of  this  discussion 
two  questions  remain  to  be  considered:  first,  that  of  heredity 
in  the  main  classes  of  tissues  of  nudticellular  organisms; 
and  second,  that  of  the  results  being  reached  by  tlie  latest 
methods  of  cytoplasmic  study  on  the  behavior  of  different 
portions  of  the  cells  in  tlie  histogenesis  of  these  tissues. 
What  is  implied  in  these  two  questions  can  be  made  clear  by 
a  special  case.  Is  the  minute  structure  of  striated  nuisele 
tissue,   for   example,    subject   to   heredity.^      If   so,    are   the 


60  The  Unity  of  the  Organism 

hereditary  attributes  determined  by  the  chromatin  or  by 
the  cytoplasm  of  the  myogenic  cells?  Applying  our  usual 
test  for  hereditary  structures,  we  ask  whether  or  not  this 
tissue  presents  attributes  characteristic  of  the  taxonomic 
groups,  species,  genera,  families  and  so  on.  Here  again, 
while  we  have  a  vast  store  of  knowledge  about  the  structure 
and  genesis  of  muscle  tissue  in  many  kinds  of  animals,  only 
incidentally,  as  a  rule,  have  the  studies  been  made  from  the 
taxonomic  standpoint. 

The  Morphology  of  Striated  Muscle  Fibers 

For  one  series  of  very  recent  studies  that  comes  near  this 
standpoint  we  are  indebted  to  H.  E.  Jordan.  On  the  taxo- 
nomic aspect  of  the  matter  Jordan  writes :  "A  quite  gen- 
eral consensus  of  opinion  considers  them  more  or  less  closely 
related,  and  ranks  them  both  between  Crustaceans  and 
Arachnoids.  Limulus  muscle,  however,  is  in  appearance  very 
much  more  like  vertebrate  than  like  insect  muscle ;  while  the 
muscle  of  the  marine  arthropod  Anoplodactylus  is  of  the 
typical  insect  type."  ^^  Of  the  numerous  differences  between 
the  fibers  of  the  two  animals  compared,  reference  to  two  will 
suffice  for  our  purpose.  They  concern  the  so-called  M 
and  Z  lines  found  in  the  light  bands  of  most  striated  muscle 
tissue.  What  Jordan  regards  as  one  of  the  important  re- 
sults of  his  work  on  Limulus  muscle  was  the  evidence  secured 
that  the  Z  line  represents  a  membrane,  as  some  observers 
have  believed,  the  specially  convincing  evidence  being  the 
fact  that  the  "line"  is  attached  to  the  sarcolemma  periph- 
erally and  to  the  nuclear  wall  centrally.  These  rela- 
tions are  lacking,  he  says,  in  the  sea-spider's  muscle.  The 
M  line,  he  tells  us,  is  especially  well  developed  and  hence 
easily  demonstrated  in  the  sea-spider  muscle  in  some  states 
of  contraction,  while  he  failed,  as  have  other  students,  to 
detect  it  at  all  in  the  Limulus  muscle. ^^ 


Evidence  from  Somatic  Histogenesis  61 

So  we  come  again  to  the  real  issue.  Assuming  the  Z 
membrane  to  be  a  cytoplasmic  structure,  as  it  has  practi- 
cally always  been  held  to  be,  are  we  going  to  deny  that  the 
cytoplasm  itself  causes  the  peculiarity  of  the  Z  membrane 
in  tlie  sea-spider  as  compared  with  that  in  Limidus,  that 
denial  being  necessitated  by  the  dogma  tliat  the  real  "seat" 
of  the  difference  is  the  chromatin  of  the  nucleus  operating 
by  some  invisible  "factor"  perhaps  of  the  nature  of  an 
enzyme? 

The  extent  of  variety  in  striated  muscle  tissue  is  brouglit 
impressively  to  view  in  such  a  comparative  study  as  that 
by  Marceau.  His  main  object  is  not  to  find  differences  but 
to  discover  whether  in  spite  of  structural  differences  they 
have  similar  traits,  as  if  they  might  all  be  derived  from  a 
single  primitive  form  which  has  undergone  more  or  less 
profound  modification. 

Of  the  many  differences  which  the  investigation  sought  to  re- 
duce to  orderliness  on  the  basis  indicated,  only  two  will  be  men- 
tioned. From  an  elaborate  table  of  measurements  of  the  diameter 
of  fibers,  we  find  the  following  results  for  the  sheep  and  pig:  "^ 

Maximum      Minimum     Average 

Sheep    25 fi  Sfx  15ju 

Pig 45  5  20 

The  other  point  selected  concerns  "striated  scleriform  trans- 
verse bands"  characteristic  of  the  muscle  fibers  of  the  vertehrate 
heart.  This  time  the  animals  we  choose  are  the  liorse  and  cow. 
The  thickness  of  the  band  is  jjiven  as  exactlv  the  same  in  thesr 
two,  but  the  distance  between  the  bands  is  1  lO^u  for  tlie  horse  and 
120/1  for  the  cow.^* 

The  FliysioJogy  of  Miiscle  Fibers 

We  might  go  on  almost  endlessly,  pointing  out  slight  hut  con- 
stant specific  differences  that  involve  differences  in  muscle  struc- 
ture, questioning  in  each  case  whether  the  hereditary  cause  of 
this  difference  lies  in  the  cytoplasm  or  chromatin  of  the  nmscle 


62  The   Unity  of  the  Organism 

cells.  But  such  repetition  would  be  useless  for  the  present  dis- 
cussion. Striated  muscle  tissue  is  specially  favorable  for  testing 
hypotheses  about  inheritance  material  from  the  functional  side 
as  well  as  from  the  structural  side.  For' example,  there  are  innu- 
merable differences,  larger  and  smaller,  in  the  limb  movements 
of  animals  belonging  to  different  species  and  genera.  I  know  of 
no  observations  which  precisely  connect  activities  of  this  sort 
with  muscle  tissue;  but  information  concerning  the  electromotor 
force  in  various  animals  is  available.  The  following  on  the  au- 
thority of  Englemann,  taken  from  Winterstein  will  serve  our 
purpose.  The  values  are  those  of  the  "demarcation  current"  of 
galvanic  electricity  of  heart  muscle,  this  current  being  generated 
by  making  a  cut  surface  at  the  base  of  the  heart  and  the  natural 
surface  at  the  apex  act  as  a  galvanic  pile:  ^^ 

Animal  species  Electromotor  force  in  D. 

Anguilla  fluviatilis 0.0265    - 

Rana  esculenta 0.031 1 

Triton  cristatus    0.0124 

Tropidonotus  natrix 0.036 

Testudo  graeca 0.022 

Columba  livia 0.0458 

Cygnus  oler   0.0168 

Mus  musculus  v.   albino 0.040 

Mus  rattus  v.  albino 0.0446 

Lepus  cuniculus 0.0o63 

These  investigations  appear  to  have  been  made  from  the  stand- 
point of  general  physiology,  and  therefore  not  to  have  been  car- 
ried out  with  the  systematic  exactness  and  exhaustiveness  de- 
manded for  taxonomic  discrimination.  We  may  consequently 
presume  that  more  searching  examination  of  the  same  series 
would  considerably  modify  these  results,  but  we  have  no  reason 
to  suppose  that  they  would  eliminate  altogether  the  differences 
due  to  the  animal  species. 

After  due  allowance  is  made  for  the  purely  physiological 
and  environic  causes  which  undoubtedly  explain  a  great 
many  of  these  differences,  probably  no  biologist  would  hesi- 
tate to  grant  that  many  of  them  have  an  hereditary  basis. 
Turning  again  to  the  question  of  the  seat  of  the  hereditary 


Evidencr  from  Somatic  Histogenesis  GiJ 

factors,  we  are  now  especially  attracted  by  the  functional 
aspect  of  the  subject.  To  an  unsophisticated  physlolog-lst 
studying  the  phenomena  involved  in  this  question,  it  would 
probably  never  occur  that  more  than  one  answer  is  possible. 
Well-informed  as  such  a  physiologist  may  be  supposed  to  hv 
on  the  important  part  known  to  be  played  by  the  nucleus  in 
the  life  of  the  cell,  he  would  undoubtedly  take  it  for  granted 
that  the  whole  nucleus,  its  chromatin  with  the  rest,  con- 
tributes in  some  fundamental  way  to  the  result.  But  unless 
well  indoctrinated  beforehand  with  the  chromosome  dogma 
of  heredity,  he  would  almost  certainly  be  amazed  were 
some  one  to  contend  seriously  that  the  cytoplasm  is  not  the 
material  basis  of  the  hereditary  peculiarities  exhibited.  He 
would  reply,  "Why,  you  are  virtually  denying  that  the 
substance  of  the  muscle  fiber  is  the  real  seat  of  muscular 
activity,  thus  implying  a  contradiction  of  the  'universally 
accepted  principle  that  the  'potential  chemical  energy  of  the 
muscle  substance  is  the  primary  source  of  muscular  energy 
in  all  its  manifestations^  ^^  for  surely  muscular  energy  *in 
all  its  manifestations'  would  include  those  elements  of  mus- 
cular acti^aty  which  are  hereditarily  distinctive  of  different 
kinds  of  animals." 

That  the  cytoplasm  is  at  least  the  main  source  of  the 
muscle  substance  furnishing  this  energy  would  not  be  ques- 
tioned, probably,  by  any  histologist,  but  the  definiteness  of 
\aew  held  at  the  present  time  on  this  subject  is  worth  re- 
calling and  is  indicated  by  such  statements  as  the  following: 
"The  energy  of  contraction  is  the  transformed  surface- 
energy  of  the  ultimate  stiiictural  elements  or  colloidal  par- 
ticles (submicrons)  composing  the  fibrils."-"^ 

Presumably  there  would  be  much  difference  of  view  among 
physiologists  as  to  the  validity  of  the  chemico-physical  part 
of  LilUe's  theory  of  muscular  contraction ;  but  apparently 
there  would  be  very  little  dissent  from  that  part  of  his  view 
which  locates  the  processes,  whatever  their  exact  nature,  in 


64  The   Unity  of  the  Organism 

the  muscle  fibers.  So  it  would  be  merely  a  matter  of  suf- 
ficient patience  to  go  over  all  the  tissue  systems,  epithelial, 
glandular,  bony,  nervous,  and  the  rest,  and  point  out  nu- 
merous certain,  and  innumerable  probable  instances  of  dif- 
ferences for  different  taxonomic  groups  of  animals,  and  to 
show  that  these  hereditary  differences  are  expressed  pri- 
marily in  the  cytoplasm  of  the  cells. 

Summary  of  Positive  Informatioji  about  the  Physical  Basis 

of  Heredity 

We  have  explored  a  vast  region  of  fact  and  theory  con- 
cerning propagation  and  development  in  organisms,  for  the 
purpose  of  ascertaining  what  is  actually  known  about  the 
organs  and  substances  by  which  hereditary  attributes  are 
produced.  Expressing  the  matter  in  terminology  familiar 
to  current  discussion  on  heredity,  we  have  been  trying  to 
find  what  is  actually  known  about  the  physical  basis  of 
heredity.  If  clear-cut,  unequivocal  information  of  the 
kind  sought  is  contained  in  all  we  have  seen,  it  ought  to  be 
statable  in  a  few  simple  sentences. 

What  has  been  accomplished  may  be  epitomized  in  two 
such  sentences : 

First.  Overwhelming  observational  evidence  has  been  se- 
cured that  the  cytoplasm  of  cells  participates  directly  in 
the  formation  of  organic  parts  which  hai^e  hereditary  at- 
tributes. 

Second.  A  great  mass  of  evidence^  partly  of  observation 
and  partly  of  legitimate  inference  from,  the  principles  of 
organic  integration,  has  been  secured,  that  the  chromosomes 
of  the  germ-cells  in  plants  and  animals  which  propagate  by 
means  of  such  cells,  participate  in  the  production  of  or- 
ganic parts  having  hereditary  attributes. 

Any  substance  which  plays  such  parts  in  development 
may  be  named  a  physical  basis  of  heredity;  and  these  two 


Evidence  from  Somatic  Histogenesis 


65 


groups,  or  categories  of  knowledge  must,  it  seems,  serve  as 
the  foundation  of  all  legitimate  reasoning  about  sucli  'i)asis 
of  heredity,"  or  "inlieritance  material,"  or  "hereditarv  fac- 
tors," or  "bearers  of  hereditary  qualities,"  or  whatever  ex- 
pression for  the  idea  be  employed. 


REFERENCE  INDEX 


1.  Payne     190 

2.  Meves     81G 

3.  Meves     817 

4.  Meves     820 

5.  Meves     850 

6.  Meves     845 

7.  Cowdry    ('14)    416 

8.  Cowdry    ('16)    436 

9.  Lewis    and   Lewis    393 

10.  Locb    ('06)     181 

11.  Conklin,    ('15)     176 

12.  Ritter    ('93) 

13.  Sedgwick     82 

14.  Minchin   ('00)    107 


15.  Goodrich     2^1- 

1 6.  Cannon    13 

17.  Cannon    23 

18.  Goebel    367 

19.  Canipl)cll     51 

20.  Wieland     197 

21.  Jordan    ('16)     493 

22.  Jordan   ('16)    fig.  7  496 

23.  Marceau    273 

24.  Marceau    280 

25.  Winterstein     Ill 

26.  Luciani    Ill,   85 

27.  Lillie,   R.    S.    ('16) 255 


Chapter  XVI . 

THE  INHERITANCE  MATERIALS  OF  GERM-CELLS 
INITIATORS  RATHER  THAN  DETERMINERS 

Antecedents  of  the  Cytoplasmic  and  Nuclear  Theories   of 

Inheritance  Material 

FOR  the  purpose  of  calling  vividly  to  mind  the  character 
of  the  evidence  on  which  the  two  propositions  rest 
with  which  the  last  chapter  ended,  it  will  be  profitable  to 
cast  a  glance  back  on  the  course  along  which  biology  has 
come  down  to  us,  with  a  view  to  finding  a  shai'ply  out- 
standing spot  in  the  early  growth  of  knowledge  which  led  to 
each  of  them.  On  the  botanical  side  such  a  spot  in  the 
knowledge  of  cytoplasm  as  hereditary  substance,  is  the 
work  of  Schleiden  on  the  microscopic  structure  of  adult 
and  developing  plant  tissues.  The  publication  of  his  Ueher 
Phytogenesis,  1838,  may  be  taken  as  the  starting  point  of 
our  knowledge  of  cellular  transformation  in  the  production 
of  tissues.  It  should  be  remembered  that  the  observers  of 
that  period  had  very  hazy  notions  about  the  distinction 
between  nucleus  and  cell-body,  or  cytoplasm.  On  the 
zoological  side  the  publication  by  Ehrenberg,  in  1836,  of 
Die  Infusionsthierchen  als  volkommene  Organismen  may,  I 
think,  be  looked  upon  as  the  first  milestone  in  the  progress 
of  knowledge  of  cytoplasmic  transformation  into  tissue  sub- 
stance. 

The  ever-broadening  stream  of  knowledge  of  the  chromo- 
somes in  relation  to  heredity  is  usually  held  to  have  orig- 
inated in   the  discovery   forty  years   ago,  by   O.   Hertwig, 

66 


Inheritance  Materials  of  Germ-Cells  61 

that  tlie  most  essential  fact  in  fertilization  is  tlie  union  of 
the  nuclei  of  the  male  and  female  germ-cells. 

That  cytoplasm  is  a  physical  basis  of  heredity  is  proved 
by  a  great  body  of  direct  observational  knowledge.  That 
the  chromatin  of  chromosomes  is  a  physical  basis  of  heredity 
is  proved  by  much  observational  knowledge  when  this  knowl- 
edge is  supplemented  by  reasoning  involving  the  principles 
of  biological  comparison  and  correlation.  These  two  masses 
of  knowledge  constitute,  as  already  indicated,  the  founda- 
tion of  all  legitimate  reasoning  about  inheritance  material. 
Whether  chromatin  and  cytoplasm  are  the  only  substances 
which  participate  in  the  formation  of  hereditary  structures 
can  not  now  be  stated  with  certainty,  though  there  are  both 
observational  and  general  grounds  for  believing  that  they 
are  not.  But  into  this  question  we  need  not  enter  in  this 
discussion.  Nor  is  it  necessary  for  our  purpose  to  inquire 
very  particularly  whether  the  conceptions  of  chromatin 
and  cytoplasm  really  imply  just  two  substances  or  two  great 
classes  of  substances,  though  it  is  best  to  have  in  mind  the 
undoubted  fact  that  the  latter  alternative  is  the  true  one. 
Beyond  a  doubt  "chromatin"  and  "cytoplasm"  ought  al- 
ways to  be  understood  to  mean  "kinds  of  chromatin"  and 
"kinds  of  cytoplasm." 

Function  of  Chromosomes  in  Heredity  Acquired  and 

Secondary 

The  question  which  specially  concerns  us  here  is  that  of 
what  the  relation  is  between  chromatin  and  cytoplasm  in 
virtue  of  which  they  play  the  particular  roles  they  are  found 
to  play  in  producing  hereditary  structures.  PiM-haps  tlie 
most  important  aspect  of  this  general  question  is  that  wliich 
the  theory  of  phyletic  evolution  naturally  brings  up:  does 
the  evidence  in  hand  suggest  any  answer  to  the  question 
whether  chromatin  or  cytoplasm  is  the  more  primitive  and 


68  The   Unity  of  the  Organism 

fundamental  as  hercditarj^  substance?  Surveying  as  we 
have  the  whole  field  of  organic  propagation,  including  the 
process  in  unicellular  organisms  as  well  as  in  multicellular, 
and  sexless  as  well  as  sexual  methods  of  reproduction,  and 
taking  the  facts  as  they  actually  present  themselves,  it 
seems  as  though  but  one  answer  to  this  inquiry  is  possible: 
The  substances  included  under  the  generic  term  cytoplasm 
are  the  more  fundamental  and  primitive. 

The  only  possible  way  of  escaping  this  conclusion  is  by 
excluding  from  the  conception  of  heredity  the  vast  majority 
of  developmental  phenomena  presented  by  unicellular  organ- 
isms and  by  monogenic  propagation  in  multicellular  organ- 
isms. As  our  examination  of  these  provinces  revealed,  such 
exclusion  is  exactly  what  the  chromatin  dogma  of  heredity 
has  undertaken,  implicitly  or  explicitly,  to  fix  upon  biology. 
The  utter  unwarrantableness  of  this  undertaking  was  made 
sufficiently  obvious,  we  may  assume,  by  the  examination ; 
so  we  need  spend  no  time  on  that  now.  All  that  is  necessary 
is  to  remind  ourselves  vividly  of  the  main  positive  outcome 
of  the  examination :  Heredity  is  a  universal  phenomenon  of 
the  living  world.  It  is  coextensive  with  organic  propagation 
and  development,  while  "carrying  heredity"  by  chromo- 
somes is,  according  to  the  evidence,  very  far  from  a  uni- 
versal phenomenon.  It  is  a  long  way  from  being  coex- 
tensive with  organic  propagation  and  development.  We 
may,  consequently,  proceed  in  our  quest  of  a  more  rational, 
more  consistent,  more  satisfactory  conception  of  the  purely 
operative  side  of  producing  hereditary  structures.  Pur- 
suing the  quest,  we  remind  ourselves  of  having  found  that 
only  the  most  meager  observational  evidence  is  afforded  by 
the  protophyta  and  protozoa,  and  by  monogenic  metaphyta 
and  metazoa,  that  chromatin  is  hereditary  substance,  while 
these  organisms  afford  an  ovei-whelming  mass  of  such  evi- 
dence that  cytoplasm  is  hereditary  substance. 

But  if  in  the  lower,  more  primitive  moiety  of  the  organic 


Tnherifance  Mafer'iah  of  Gcrm-CcUs  69 

realm,  clironiatln  is  a  ])lijsic'al  basis  of  litTcdity  to  only  a 
limited  extent  and  in  a  partial  way  but  has  this  office  widely 
and  positively  fixed  in  the  higher  moiety,  the  moiety,  tliat  is, 
in  which  bisexual  propagation  is  I'ully  establislied,  uliat 
other  conclusion  can  be  drawn  consistently  Avith  the  modes 
of  reasoning  universally  sanctioned  by  evolutionists,  than 
that  the  function  of  "carrying  hereditary  qualities"  by  the 
chromosomes  in  higher  organisms  is  a  secondary  or  ac- 
quired, or  better  a  delegated  or  assigned  function?  Hered- 
ity is  far  older,  phylogenetically,  and  far  broader  tax- 
onomically  than  is  the  chromatin  mechanism  by  which  it 
now  in  part  manifests  itself.  Under  this  interpretation  the 
acquisition  by  chromosomes  of  the  function  of  carrying 
heredity  would  belong  to  the  same  evolutional  type  as  for 
example  the  acquisition  by  certain  cells  of  the  function  of 
muscular  contraction,  or  by  certain  other  cells  of  con- 
ducting nervous  stimuli.  The  advantage  and  satisfaction 
of  a  conception  of  the  role  of  chromosomes  in  heredity 
which  ranges  them  naturally  and  easily  with  all  other  or- 
gans and  tissues  of  the  plant  and  animal  body  will  be 
quickly  seen  by  every  one  to  whom  the  seemingly  endless 
chance  of  discovering  new  interrelationships  and  consis- 
tencies jn  living  nature  is  one  of  the  most  rewarding  things 
about  biological  investigation. 

While  we  are  duly  impressed  with  the  importance  of  per- 
ceiving that  chromosomes  fall  into  the  great  class  of  otlier 
organs  and  tissues  when  considered  from  the  standpoint 
of  phyletic  differentiation  of  structure  and  function,  we 
should  not  fail  to  notice  that  within  the  class  they  hold  on 
a  number  of  counts  a  very  distinct  place.  Probably  the 
most  distinctive  of  these  counts,  at  any  rate  the  one  most 
important  for  this  discussion,  is  the  fact  that  while  the  vast 
majority  of  tissues,  taking  the  term  in  its  usual  meaning, 
stand  at  or  near  the  termination  of  the  ontogenic  series — 
are,  in  other  words,  the  final  stage  in  the  series — the  chro- 


70 


The   Unity  of  the  Organism 


mosomes  in  their  function  as  bearers  of  heredity  stand  at 
the  beginning.  They  represent  the  initial  stage  in  the  series. 
Furthermore,  their  function  in  this  respect  is  unique  as 
contrasted  with  the  function  of  other  tissues,  in  that  while 
other  tissues  liave,  typically,  each  a  single  function  wliich 
they  perform  immediately,  the  chromatin  of  a  given  germ- 
cell  has  a  great  complex  of  functions,  namely,  that  of  ini- 
tiating the  development  through  which  all  the  attributes  of 
the  individual  during  its  whole  life-career  are  developed. 
In  a  word,  chromosomes  of  germ-cells  are  not  determiners  or 
carriers  of  determiners ;  they  are  initiators  or  carriers  of 
initiators.  They  may,  then,  be  called  bearers,  or  carriers 
of  heredity  in  a  very  literal  sense,  namely  in  the  sense  that 
they  are  made  use  of  by  one  individual,  the  parent,  to  carry 
across  or  transfer  from  itself  to  another  individual,  the 
child,  the  hereditary  attributes  of  tlie  species  in  a  latent  or 
potential  state.  By  virtue  of  their  being  thus  used,  they 
are  members  of  a  developmental  series,  in  which  series  their 
place  is  at  the  beginning  and  not  at  the  end,  the  nature  of 
the  series  depending  on  the  phylogenic  history  of  the  par- 
ticular organism  to  which  the  particular  chromosomes  be- 
long. 

The    Two- fold   Character    of    the   Problem    of   Hereditary 

Substance 


At  this  point  it  becomes  a  matter  of  the  greatest  im- 
portance for  theories  about  hereditary  substance  to  dis- 
tinguish between  the  problem  of  the  operation  of  such  sub- 
stance in  the  developing  individual,  and  that  of  how  sucli 
substance  ever  came  to*  be  hereditary  substance;  stated 
otherwise,  between  the  problems  of  how  any  substance  par- 
ticipates either  directly  or  as  an  agent  in  the  building  up  of 
a  structure  having  hereditary  attributes,  and  that  of  how 
the  substance  itself  became  impressed  with  the  attributes,  in 


Inheritance  Materials  of  Germ-Cells  71 

a  latent  state,  of  tlie  progenitors  of  the  developing  in- 
dividual. 

Investigation  of  the  first  of  these  j)roblems  is  to  a  large 
extent  a  matter  of  observation,  as  we  have  seen  in  the  pre- 
ceding pages.  The  sub-science  of  histogenesis  consists 
largely  in  tracing  out  the  processes  by  which  completed 
tissues  arise  by  the  transformation  of  less  differentiated  or 
undifferentiated  cells.  And  when  such  newly  arisen  tissues 
and  structures  are  proved  to  be  hereditary  by  such  evi- 
dence as  we  have  called  attention  to,  then  the  study  of 
histogenesis  comes  to  be  so  far  a  study  of  hereditary  sub- 
stance. 

When,  however,  we  turn  to  the  other  problem,  that  of 
how  hereditary  substance  comes  to  be  such,  we  are  in  a 
different,  a  much  more  difficult  case,  for  so  far  science  has 
succeeded  in  getting  almost  no  obsei'vational  hold  upon  it; 
and  despite  the  vast  discussion  it  has  received  the  darkness 
that  envelops  it  is  hardly  an  iota  less  black  than  it  was  the 
day  of  its  original  formulation.  But  stygian  as  the  darkness 
is,  here,  especially  as  to  details,  we  yet  are  able  to  see, 
probably,  the  quarter  from  which  light  will  come  if  ever  it 
does  come.  That  quarter  is  the  physical-chemistry  con- 
ception of  the  organism  as  a  system  of  phases  the  whole  of 
which,  as  a  species  entity,  is  essential  to  its  equilibrated 
activities.  This  nature  of  the  organism,  together  with  some- 
thing akin  to  its  internal  secretory  and  enzymic  productiv- 
ity, enables  it,  we  may  conjecture,  by  some  means  now 
wholly  unknown,  to  reflect  its  totality  of  transferable  at- 
tributes upon  the  germinal  cells  and  to  transform  thrni  into  a 
latent  state.  Ungrudging  acknowledgment  of  the  complete- 
ness of  our  ignorance  of  how  any  part  of  a  cell  or  any  other 
portion  of  an  organism  becomes  endowed  witli  the  capacity  to 
develop  or  causally  to  affect  the  development  of  an  organism 
similar  to  that  from  which  it  came,  should  he  an  important 
item  in  the  preparation  to  accept  any   and   all  indubitable 


72  The   Unity  of  the  Organism 

facts  connected  with  heredity,  even  though  no  causal  ex- 
planation of  them  is   forthcoming. 

The  childlikeness,  as  Conklin  well  characterized  it,  of 
the  belief  that  chromosomes  are  a  simple  and  complete  ex- 
planation of  inlieritance  would  not  be  so  bad  in  itself.  If 
it  stopped  there,  as  genuine  childHkeness  would,  no  posi- 
tive harm  could  be  done.  It  is  the  making  of  this  belief 
the  starting  point  of  a  grand  speculation  which  blinds  the 
eyes  and  closes  the  mind  to  a  vast  number  of  facts  and 
legitimate  inferences  about  heredity,  that  plays  havoc  with 
thinking  on  genetics.  It  would  be  a  great  gain  if  gen- 
etic theory  would  recognize  wholeheartedly  that  all  or^ 
ganic  development,  as  contrasted  with  mere  enlargement, 
consists  more  fundamentally  and  obviously  in  transforma- 
tion of  substances  than  it  does  in  unchanged  continuity  of 
substances.  For  under  such  recognition  the  futility  of 
attempting  to  explain  the  transfonnation  of  one  lot  of 
substance  by  referring  it  to  another  lot  which  does  not 
transform,  or  in  other  words  to  explain  development  by 
something  that  does  not  itself  develop,  would  be  manifest. 

That  the  chromosome  theory  of  heredity  in  reality  deep- 
ens rather  than  illumines  the  darkness  which  surrounds  the 
problem  is  seen  when  one  reflects  that  not  only  does  it 
throw  no  light  on  the  question  of  how  the  chromosomes  come 
to  be  bearers  of  heredity,  but  that  it  creates  the  new  and 
equally  difficult  question  of  how  the  chromosomes  (which  ac- 
cording to  the  theory  maintain  unchanged  their  individuality 
not  only  from  generation  to  generation  but  throughout  each 
ontogeny)  are  yet  able  to  be  causally  operative  in  the  cell- 
bodies  undergoing  the  transformations  which  they  actually 
do  undergo  in  the  developing  organism.  That  the  germ- 
plasm-chromosome  theory  of  heredity  could  have  led  its 
devotees  to  sidestep  the  details  of  ontogeny,  especially  those 
of  histogenesis,  to  the  extent  which  our  review  has  shown 
it    to    have   done,    would   be    unbelievable   but   for   what    is 


Inheritance  Materials  of  Germ-CeUs  73 

actually  before  us  in  tlie  recent  history  of  hioloirv. 

Aiewin^tp  heredity  as  bein^-  dcfinitiyely  a  kind  of  organic 
transformation — transformation,  that  is,  in  accorchmce  witli 
a  pre-existing  or  ancestral  ])attern  more  than  it.  is  a  kind 
of  continuity — it  becomes  obyious  that  even  were  the  dem- 
onstration to  become  complete  that  the  chromosomes  arc 
the  only  ])ortions  of  the  germ-cells  *  essential  to  fertiliza- 
tion, they  still  would  not  be  proved  bearers  of  heredity  in 
such  sense  as  the  germ-plasm  theory  holds  them  to  be.  They 
would  not  because  the  problem  of  the  transformations 
which  constitute  ontogeny  would  still  be  initouched.  The 
theory  would  be  established  only  when  the  demonstration 
should  be  produced  that  the  chromosomes  cause  immedi- 
ately all  the  particular  ontogenic  transformations  known 
to  be  hereditary.  All  that  would  be  proved  about  heredity 
by  demonstration  that  the  chromosomes  alone  participate 
in  fertilization  would  be  that  the  chromosomes  alone  con- 
stitute the  first  ontogenic  stage  of  the  hereditary  parts  of 
the  particular  organism  to  which  the  fertilized  Qgg  gives 
rise. 

The  Frohahility  That  Inheritance  Material  Becomes  Siicli 

In  Each   Ontogeny 

But  because  thus  far  failure  has  attended  all  efforts  to 
get  knowledge  of  how  hereditary  substance  is  produced, 
are  we  obliged  to  own  that  we  know  nothing  at  all,  even 
inferentially,  about  its  production?  And  is  tlie  search  for 
such  knowledge  to  be  given  up  as  hopeless?  ^fy  answer  is 
an  energetic  negative  to  both  these  (juestions.  In  tlie  first 
])lace,  there  is  much  evidence  to  support  the  hypothesis, 
very  general  to  be  sure  but  yet  by  no  means  devoid  of  use- 
fulness,    that    hereditary    substance    becomes    such    in    some 

*  The  utter  uinvarrantableiiess  of  the  eonmion  assumption  that  as 
regards  the  male  germ-eell  sueh  a  demonstration  is  "praetieally  eom- 
plete"  will  be  noticed  presently. 


74  The  Unity  of  the  Organism 

way  through  being  subject  to  the  metabolic  processes  com- 
mon to  the  whole  organism.  Undoubtedly  the  germ-plasm 
dogma  itself  has  tended  strongly  to  divert  attention  from 
this  aspect  of  the  problem  of  germinal  material — indeed, 
has  tended  to  minimize  the  importance  of  the  metabolism 
of  such  material  even  if  it  has  not  tended  to  deny  that  the 
material  is  subject  to  this  process. 

So  important  is  it  from  the  organismal  standpoint  to 
conceive  the  material  basis  of  heredity  as  part  and  parcel 
of  the  organism  generally,  especially  as  regards  the  basal 
growth  and  sustentative  processes,  that  we  must  examine  in 
some  fullness  the  evidence  favorable  to  such  a  conception. 
In  its  most  brazenly  evidence-ignoring  form,  the  gerai-plasm 
dogma  asserts  that  the  female  parent  does  not  really  pro- 
duce the  eggs  or  the  male  parent  the  sperm,  as  they  seem 
to,  but  that  these  are  produced  by  previous  germs  ad  in- 
finitum. There  are,  to  be  sure,  quite  a  number  of  observ- 
able facts,  as  those  of  the  early  formation  of  germ-cells 
in  several  animals,  that  can  be  forced  into  a  seeming  sup- 
port of  such  a  conception.  But  the  familiar  and  all  but  uni- 
versal fact  that  multicellular  organisms,  plants  and  animals 
alike,  are  sexually  immature  for  a  shorter  or  longer  part  of 
their  lives,  the  very  essence  of  the  immaturity  being  the  un- 
developed state  of  the  reproductive  system,  would  be  a 
sufficient  refutation  of  the  view  for  any  mind  not  made 
impervious  to  facts  by  long  and  faithful  sophistication. 

Germ-Cells  Subject  to  Metabolism  Like  All  Other  Cells 

The  biological  commonplace  that  all  germ-cells,  like  all 
other  cells,  undergo  a  process  of  growing  and  maturing 
before  they  can  perform  their  distinctive  office,  and  that 
this  process  depends  upon  the  retention  of  the  germs  by  the 
parent  organism,  ought,  as  already  indicated,  to  be  a  suf- 
ficient   antidote    againjst    the    germinal    continuity    fallacy, 


Inheritance  Materials  of  Germ-CeUs  75 

even  tliough  nothing"  were  known  as  to  exactly  what  goes 
on  in  the  germ  while  it  is  growing  and  ripening.  Ihit  we 
are  by  no  means  without  positive  knowledge  uiuUr  this 
head.  In  fact  the  last  few  cell  divisions  innnediately  pre- 
ceding the  ripening  of  both  ova  and  spermatozoa,  and  the 
ripening  processes  themselves,  have  received  searching  ex- 
amination during  tlie  last  few  decades,  with  the  result  that 
hardly  any  cytological  phenomena  are  better  known  than 
are  the  profound  morphological  changes  which  accompany 
these  j)rocesses.  That  these  changes  are  particularly  mani- 
fest in  the  chromosomes,  the  assumed  seat  of  the  determiners 
of  heredity,  is  one  of  the  very  things  that  has  aroused  so 
much  interest  in  the  processes.  Nor  are  we  wholly  unin- 
formed about  the  chemical  changes  taking  place  in  the 
growing  germ-cells.  Unfortunately  knowledge  in  this  field 
has  hardly  passed  the  stage  of  early  infancy,  but  at  least 
enough  is  known  to  warrant  the  assertion  that  the  young 
germ-cells  are  subject,  as  are  all  the  other  cells,  to  the 
general  metabolism  of  the  organism. 

Chemical  Changes  in  Germ-Cells  During  Parcnfs  Ontogcnjj 

About  the  most  striking  information  wc  liave  in  this 
field  is  what  has  come  from  such  investigations  as  those 
on  the  chemical  changes  which  occur  in  the  sex  glands  and 
other  body  parts  during  reproduction  in  some  fishes.*  Mic- 
scher's  work  was  ground-breaking  in  this  domain  for  it 
was  the  first  to  show  that  the  "sexual  organs  in  the  salmon 
develop  at  the  expense  of  the  muscular  system,  and  tliat  the 
salmine  deposited  in  the  testis  during  the  breeding  season 
must  be  derived  from  the  proteins  of  the  nuiscle,  since  the 

*  Notable  among  these  studies  are:  Histocliemische  und  physiolotjischr 
Arbeiten,  gesammelt  und  herausf/ff/ehev  von  seinen  Freiindin,  by  Mio- 
cher;  and  Changes  in  the  Chemical  Composition  of  the  Ilerrina  dtirina 
the  Reproduction  Period,  by  Milroy.  Biochemical  Journ.,  v.  iii,  1908,  p. 
366. 


76  The  Unity  of  the  Organism 

fish  does  not  take  any  food  during  the  period."  ^  The  work 
of  Riddle  and  his  collaborators  is  producing  evidence  to  the 
same  effect. 

Such  researches  do  not,  to  be  sure,  prove  that  the  chem- 
ico-physiological  changes  extend  to  the  chromosomes  of  the 
germ-cells,  much  less  to  the  imaginary  determiners  of  hered- 
ity in  the  chromosomes.  But  viewing  the  results  in  the 
light  of  the  well-grounded  general  belief  that  the  mdst 
fundamental  test  of  living  substance  is  metabolic  change, 
it  is  seen  that  any  hypothesis  which  assumes  the  existence 
in  the  germ-cells  of  something  virtually  not  subject  to  the 
general  metabolism  of  the  cells,  assumes  at  the  same  time  the 
burden  of  furnishing  objective  evidence  that  such  a  something 
does  exist. 

As  a  matter  of  fact  the  prime  offense  of  the  gerai-plasm- 
determiner  hypothesis  is  that  its  very  essence  places  it 
beyond  the  reach  of  scientific  observation.  Such  truth  as  it 
may  contain  cannot  be  made  really  effective  because  it  can 
not  be  proved,  and  such  error  as  it  may  contain  can  not 
be  robbed  of  its  power  for  evil  because  it  cannot  be  dis- 
proved. In  a  word,  the  hypothesis  is  one  that  belongs  to 
the  realm  of  dialectics  primarily,  and  has  no  just  claim  to 
a  place  in  inductive  science. 

The  Possibility  of  Changing  Sex  By  Influences  on  the  Germ 

But  perhaps  the  most  conclusive  evidence  of  the  funda- 
mental dependence  of  true  germinal  material  upon  the  or- 
ganism, should  somewhat  fuller  verification  of  the  obser- 
vations be  obtained,  are  results  like  those  reached  by  Whit- 
man, King,  Whitney,  Riddle,  and  by  R.  Hertwig  and  his 
students,  according  to  which  sex  may  be  reversed  in  several 
animal  species  by  various  conditions  extraneous  to  the 
germ  itself,  acting  on  the  germ-cells  from  which  the  animal 
is  to  develop.  The  instance  of  this  usually  regarded  as  best 
established  is  afforded  by  certain  species  of  frogs  and  toads. 


Inheritance  Materials  of  (icrvi-Cclls  77 

The  widely  known  result  reached  hy  Hertwig  and  vurlHed 
and  extended  by  Kuschakewltsch  ex})re.ssed  in  a  single  sen- 
tence, is  that  the  number  of  males  and  of  females  ])ro(luced 
by  the  eggs  of  a  given  female  depends  upon  whether  the 
eggs  are  fertilized  when  newly  rij)e  or  when  over-ri})e,  a  great 
])redominance  of  males  coming  from  the  latter  class.  In 
a  tabulation  of  the  results  of  four  sets  of  experiments  \)\ 
Hertwig "  almost  every  case  shows  the  number  of  males 
increased  when  the  time  elapsing  })etween  deposition  of  the 
eggs  and  fertilization  was  increased,  the  highest  percentage 
of  males  in  any  one  lot  being  759,  fertilization  in  this  in- 
stance having  been  twenty-two  hours  after  the  fertilization 
of  the  last  preceding  lot. 

In  a  species  of  toad,  Bufo  lentigijwsus,  results  have  been 
obtained  just  the  reverse  of  those  on  the  frog,  that  is,  the 
proportional  number  of  females  has  been  experimentally 
increased.  This  was  accomplished  by  fertilizing  the  eggs 
in  water  made  slightly  alkaline.  Since  frog  eggs  are  known 
to  absorb  water  when  they  remain  long  in  it,  as  in  the  case 
of  those  which  gave  rise  to  a  preponderance  of  males  in 
the  Hertwig  method  of  experimenting;  and  since  alkaline 
solutions  extract  water  from  eggs,  and  likewise  cause  them 
to  produce  a  preponderance  of  females,  Miss  King  drew 
the  obvious  conclusion  that  the  quantity  of  water  contained 
in  eggs  of  these  animals  may  be  a  factor  in  determining  the 
sex  of  the  animals  developed  from  the  eggs.  Although 
Miss  King  recognizes  that  her  experiments  do  not  furnish 
final  proof  of  the  conclusion  she  draws,  she  believes,  rightly 
it  would  seem,  that  they  weigh  heavily  in  tliat  direction. 
"As  they  stand,"  she  writes,  "the  results  strongly  suggest 
that  sex  in  Bufo  is  determined  at  or  near  the  time  of  fer- 
tilization, and  that  external  factors  acting  during  this  ])o- 
riod  may  influence  the  sex-determining  mechanism  in  such 
a  way  as  to  cause  it  to  produce  one  sex  or  the  other.  The 
results  also  seem  to  indicate  that  in  Ilufo  sex  is  determinwl  in 


78  The  Unity  of  the  Organism 

the  ^ggi  and  that  it  may  depend  in  some  way  on  the  rela- 
tive amount  of  water  in  the  ^gg  at  the  time  of  fertiliza- 
tion." 3 

Riddle,  perhaps  the  most  outspoken  opponent  of  sex  pre- 
destination now  writing,  strongly  espouses  the  hypothesis 
that  the  sex  to  which  a  particular  Qgg  will  give  rise  is  de- 
pendent partly  on  the  quantity  of  water  which  that  Qgg 
contains.  But  whether  water  is  a  factor  in  determining  sex 
or  not,  the  evidence  presented  by  Riddle,  coming  partly  from 
researches  by  C.  O.  Whitman  and  partly  from  his  own, 
constitutes,  when  taken  with  the  evidence  to  the  same  effect 
presented  by  other  investigators,  almost  if  not  quite  com- 
plete proof  that  sex  is  not  the  hard-and-fast  thing  which 
most  present-day  genetic  speculation  would  make  it. 

Furthermore  the  evidence  produced  by  these  two  inves- 
tigators seems  to  connect  the  decision  as  to  which  sex  a 
particular  ^gg  shall  give  rise,  with  some  condition  of  the 
parents.  It  is  well  known  to  all  zoologists,  in  the  United 
States  at  least,  that  at  the  time  of  his  death  Professor  Whit- 
man had  accumulated  a  vast  store  of  data  on  the  habits, 
particularl}'^  the  breeding  habits,  of  pigeons.  To  Doctor 
Riddle,  who  had  worked  with  Whitman  considerably,  fell 
the  task  of  carrying  on  to  some  extent  Whitman's  experi- 
ments and  of  preparing  for  publication  the  results  which 
Whitman  left  in  the  rough.  The  following  quotation  from 
Riddle's  paper  referred  to  above,  summarizes  Whitman's 
results  that  are  especially  important  for  us  now:  "Whit- 
man found  that  if  certain  very  distantly  related  pigeons 
[i.e.,  two  individuals  from  different  families]  are  mated 
that  only  male  offspring  resulted.  If  the  matings  were 
made  of  individuals  not  quite  so  distantly  related — different 
genera  usually — and  if  to  this  situation  be  added  the  ele- 
ment of  overwork  at  reproduction  [i.e.,  the  birds  not  being 
permitted  to  nest  their  own  eggs,  but  forced  to  keep  laying 
eggs   in   rapid   succession]    then   the   first   several  pairs    of 


Inheritance  Materials  of  Germ-Cells  79 

eggs  produced  in  the  spring  will  produce  all  or  nearly  all 
males.  The  last  several  pairs  of  eggs  laid  in  autumn  will 
produce  all,  or  nearly  all,  females.  At  the  transition  period 
in  the  summer  he  found  that  some  pairs,  or  clutches,  of  eggs 
produced  both  a  male  and  a  female.  In  these  cases  it  was 
usually  the  first  Qgg  that  produced  the  male;  and  the  sec- 
ond e^gg — laid  forty  hours  after  the  first — that  gave  rise 
to  a  female."  ^ 

Into  Riddle's  interesting  discussion  of  Whitman's  results 
and  his  own  chemical  studies  on  the  eggs  of  pigeons  and 
hens  we  need  not  go.  Suffice  it  to  say  that  it  seems  to  me 
Riddle  is  justified  by  the  evidence  now  in  our  possession, 
in  his  contention  that  "sex  rests  upon  a  quantitative  and 
reversible  basis"  and  that  in  this  sense  it  has  been  controlled 
by  conditions  extraneous  to  the  genn-cells  themselves.  This 
does  not  imply,  as  I  understand,  that  such  control  would 
necessarily  be  practicable  of  even  possible  in  all  organisms, 
nor  does  it  preclude  the  possibility  that  in  some  species 
there  may  be  dimorphic  or  partially  dimorphic  spermatozoa 
or  ova  as  regards  sex  production.  Neither  does  it  preclude 
the  possibility  that  in  some  cases  where  a  preponderance 
of  one  sex  has  been  observed,  this  is  due  to  selective  mor- 
tality or  some  process  other  than  the  actual  shifting  of  the 
sex  tendency  in  the  particular  eggs. 

These  several  concordant  bodies  of  testimony  must,  it 
would  appear,  open  the  eyes  of  biologists  sooner  or  later  to 
the  ludicrousness  of  a  theory  that  would  make  the  parent 
organism  hardly  more  than  a  combined  culture  medium  and 
incubating  oven  for  its  germ-cells. 

The  Determiner  Conception  Contrary   to   Ordinarif   Chem- 
ical Principles 

If,  on  the  basis  of  such  facts  as  we  have,  we  try  to  come 
still  closer  to  the  questions  of  how  the  assimilative  and 
morphogenic  processes   of  the   organism   occur,  whether  in 


80  The  Unity  of  the  Organism 

the  production  of  hereditary  substance  or  in  the  transform- 
ation of  such  substance  into  actual  hereditary  structures 
and  activities,  we  find  ourselves  hedged  about  on  every  side 
by  partial  knowledge,  by  dubious  knowledge,  and  by  com- 
plete ignorance  on  many  fundamental  points. 

However,  chemical  considerations  seem  to  point  the  way 
to  future  discovery.  In  the  first  place,  it  seems  necessary 
to  recognize  that  the  whole  germ-plasm  conception  as  orig- 
inally promulgated,  with  its  interminable  system  of  "bear- 
ers" and  consummators,  was  contrary  to  what  is  well  known 
about  chemical  processes  generally.  Thus  the  continuity 
presented  by  a  complex  chemical  operation  does  not  consist 
in  an  unchanged  series  of  individual  entities  of  some  sort, 
such  as  determinants  and  determiners  are,  or  originally 
were  conceived  to  be,  but  rather  in  a  regular  succession  of 
transformations.  For  example,  when  chromic  hydroxide, 
which  is  grayish-green,  is  dissolved  in  acid,  a  green  solu- 
tion results,  which  turns  to  greenish  violet  or  pure  violet 
if  allowed  to  stand  a  long  time.  Exactly  what  the  chemical 
changes  are  that  correspond  to  these  color  changes  I  do 
not  know,  and  probably  the  information  which  chemists  have 
on  the  subject  is  not  exhaustive.  At  any  rate  modem 
chemistry  conceives  the  phenomenon  to  consist  in  a  succes- 
sion of  reactions  and  transformations,  the  various  colors 
and  shades  being  attributes  of  the  compounds  that  exist 
in  the  various  stages  along  the  way,  and  not  as  due  to  in- 
dividual bodies  carried  by  the  preceding  stages  for  the 
express  and  exclusive  purpose  of  producing  the  particular 
colors  that  are  observed,  as  would  be  implied  in  such  a 
metaphysical  scheme  as  was  the  germ-plasm  theory  elab- 
orated by  Weismann. 

Enzymic  chemical  action  presents  perhaps  a  still  better 
starting  point  for  imagining  what  the  fundamental  hered- 
itary processes  may  be  than  does  ordinary  chemical  activity 
like  that  just  instanced.     The  essence  of  this  kind  of  activ- 


Inheritance  Materials  of  Germ-Cells  81 

ity  is,  as  everybody  knows,  that  in  some  way  enzymes  cause 
or  at  least  facilitate  transformation  in  other  substances. 
Thus  the  attribute  of  solubility  of  tlic  sugar  Into  whicli 
starch  is  transformed,  through  tlie  action  of  tlie  saHvarv 
enzyme  ptyalin,  is  not  held  to  be  due  to  a  determiner  f(}r 
solubility  carried  by  the  ptyalin  and  passed  on  into  the 
sugar,  but  rather  it  is  recognized  tliat  sohibiHty  is  one  of 
the  attributes  possessed  by  the  kind  of  sugar  into  ^vhich 
starch  is  converted  by  the  ptyalin.  The  solubility  is  thought 
of  rather  as  an  attribute  of  the  sugar  and  not  as  something 
once  latent  in  the  ptyalin  which  produced  the  sugar.  A 
few  details  of  the  action  of  the  enzj^me  in  this  case  illustrate 
the  point  still  better.  Maltose,  which  is  the  chief  if  not 
the  only  sugar  resulting  from  the  action  of  ptyalin,  is  not 
reached  by  a  single  bound,  as  one  might  say,  but  through  a 
series  of  bodies  known  as  dextrins,  at  least  three  of  wliich 
have  been  recognized.  These  are  amylo-  erythro-  and 
achroo-dextrin,  named  from  the  color  they  display  Avhen 
treated  with  iodine,  the  first  mentioned  turning  blue,  tlie 
second  red,  and  the  third  remaining  colorless.  What  mod- 
ern chemist  would  think  of  explaining  the  blue  of  the  amylo- 
dextrin  by  a  "determiner"  for  that  color  in  the  ptyalin 
or  even  in  the  starch,  the  red  of  the  erythrodextrin  by  an- 
other determiner  for  red,  and  so  on.?  That  is  the  sort  of 
explaining  chemists  of  a  century  ago  did,  but  they  have 
long  since  learned  not  merely  the  futility  but  the  scientific 
evil   of  such  explanation. 

If  it  were  germane  to  our  ])resent  task  we  might  go  on 
and  show  that  the  gene  conception  in  modern  giMietics  is 
really  a  revival  in  biology  to-day  of  the  geri-e  conception 
which  passed  muster  in  chemistry  a  hundred  years  ago, 
when  oxygen  and  hydro(7r7J  were  named.  Such  an  exposi- 
tion would  be  appropriate  to  a  history  of  scientific  theory 
or  to  a  treatise  on  the  theory  of  natural  knowledge,  hut 
hardly  to  the  present  work. 


82  The  Unify  of  the  Organism 

Endorsement   of  E.   B.    Wilson's   Proposal   to   Drop  "De- 
termvner^'  From  the  V ocahulary  of  Genetics 

In  his  Croonian  Lecture  having  the  title  The  Bearing  of 
Cytological  Research  on  Heredity,  E.  B.  Wilson  said,  "In 
the  meantime  it  would  be  well  to  drop  the  term  'determiner' 
or  'determining  factor'  from  the  vocabulary  of  both  cytol- 
ogy and  genetics."  ^  If  the  facts  and  arguments  set  forth 
in  the  preceding  pages  are  valid,  they  constitute  a  demon- 
stration that  not  only  would  it  "be  well  to  drop  the  term 
'determiner,'  "  but  that  it  must  be  dropped,  at  least  in  its 
present  application,  before  thought  and  investigation  on  the 
mechanism  of  heredity  can  be  free  and  in  very  deed  truth- 
seeking.  "What  we  reall}'^  mean  to  say,"  Wilson  continues, 
"is  'differential'  or  'differential  factor,'  for  it  has  become 
entirely  clear  that  every  so-called  unit  character  is  pro- 
duced by  the  cooperation  of  a  multitude  of  deteiTnining 
causes."  So  far  as  these  statements  go  they  are  in  strict 
accord  with  the  organismal  standpoint  maintained  in  this 
volume,  and  we  may  also  say,  with  the  physical-chemistry 
standpoint. 

Where  attributes  of  adult  organisms  have  been  so  defin- 
itely correlated  with  particular  chromosomes  and  possibly 
parts  of  chromosomes  of  the  germ-cells  as  seems  to  be  the 
case  in  the  fruit  flies,  such  chromosomes  are  unquestionably 
differential,  and  since  they  stand  at  the  very  beginning  of  a 
long  and  complex  transforming  and  developing  series,  they 
may  very  properly  be  called  differential  factors  even  though 
they  do  not  themselves  participate  substantively  in  the 
transformation.  The  general  similarity  of  their  mode  of 
action  to  that  of  enzymes  is  certainly  considerable :  a  minute 
quantity  of  the  substance  is  capable  of  inducing  or  facili- 
tating the  transformation  of  a  large  amount  of  other  sub- 
stance in  a  perfectly  definite  manner,  and  the  inducing  agent 
is  not  itself  consumed. 


Inheritance  Materials  of  Germ-Cells  83 

Advantages  of  Conceiving  Germ-Cell  Chromosomes  as  Initia- 
tors in  Hereditary/  Development 

This  chemical  way  of  viewing  chromatin  and  chromosomes 
sanctions  the  idea  that  these  are  to  be  regarded  as  initiators 
of  developmental  processes  which  lead  to  hereditary  attri- 
butes, rather  than  determiners  of  those  attributes.  If  one 
wants  to  know  in  what  way  this  conception  would  liave 
an  advantage  over  the  determiner  conception  as  a  working 
hypothesis,  my  reply  is  that  the  advantage  is  two-fold. 
First,  it  would  surely  correct  the  tendency  of  genetic  re- 
search under  the  guidance  of  the  determiner  hypothesis,  to 
restrict  its  attention  to  attributes  of  adults  at  one  end  of 
the  ontogenic  series  and  to  the  chromosomes  of  the  germ- 
cells  at  the  other  end,  and  to  ignore  or  touch  only  in  the 
lightest  way  all  the  intervening  parts  of  the  series.  This 
correction  would  result  because  the  new  standpoint  would 
bring  the  whole  series  of  continuities  and  transformations 
alike  into  proper  perspective,  revealing  thus  that  the  mem- 
bers of  the  series  intervening  between  germ  and  adult  must 
be  investigated  in  exactly  the  same  way  and  with  the  same 
objects  in  view  as  the  end  members,  if  complete  understand- 
ing of  the  hereditary  process  be  the  goal  of  research.  It 
could  not  then  happen  that  the  egg-cell  would  be  repre- 
sented, as  it  now  so  commonly  is,  as  a  relatively  large  sac 
containing  nothing  significant  for  heredity  except  the  rela- 
tively small  chromosomes.  Nor  could  nearly  the  whole 
mass  of  ontogenic  phenomena,  especially  those  of  histo- 
genesis, be  treated  so  lightly  in  speculation  and  so  largely 
neglected  in  investigation  as  they  have  })een  under  the 
domination  of  the  determiner  theory. 

The  second  advantage  in  the  initiator  conception  is  tliat 
since  it  would  recognize  the  "differential  factor"  of  tin- 
chromosomes  to  be  in  reality  due  to  the  fact  that  the  wliole 
ontogenic  series  to  which  the  chromosomes  belong  is  dilfLr- 


84  The   Unity  of  the  Organism 

ential,  that  is,  that  it  pertains  to  a  particular  species  or 
kind  of  organism,  it  would  put  an  end  to  the  notion  by 
which  recent  genetic  science  has  been  so  largely  dominated, 
that  the  problem  of  how  the  series  came  to  be  thus  specific 
or  differential  may  be  solved  by  speculation,  and  it  would 
incite  geneticists  to  efforts  to  solve  the  problem  by  obser- 
vation aided  by  experiment.  It  is  impossible  to  refute  the 
charge  that  genetics  is  to-day  more  interested  in  an  elab- 
orate system  of  conceptions — of  speculation,  in  other  words 
— than  it  is  in  obsei'ved  or  possibly  observable  phenomena. 
We  cannot  keep  too  constantly  before  our  minds  the  fact 
of  our  almost  complete  ignorance  of  how  any  substance 
becomes  hereditary  substance  whether  through  the  "inheri- 
tance of  acquired  characters"  or  in  any  other  way.  Hence 
mere  speculation  on  the  subject  after  the  manner  of  the  pan- 
gens  idea  is  much  worse  than  nothing  if  permitted  to  run  into 
a  bewildering  and  enslaving  system  like  that  of  the  germ- 
plasm  theory  as  it  came  from  Weismann's  mind.  Neverthe- 
less it  is  quite  germane  to  the  present  discussion  to  point 
out  that  whatever  might  be  the  nature  of  the  chemical  ac- 
tion, whether  enzymic  or  some  other,  through  which  the 
series  of  ontogenic  transformations  should  be  accomplished, 
the  character  and  subtlety  of  these  processes  seem  to  make 
them,  more  than  any  others  we  know,  competent  with  some 
modification  to  serve  as  the  go-between  for  impressing  the 
germinar material  with  the  latent  attributes  of  the  species. 

Inconclusiveness  of  the  Cytological  Evidence   Usually  Ap- 
pealed to  in  Support  of  the  Chromosome  Theory 

And  this  leads  to  the  concluding  statements  of  this  dis- 
cussion. The  three  categories  of  cytological  fact  which 
have  been  weightiest  in  the  formation  and  maintenance  of 
the  chromosome  theory  of  heredity  are  the  individuality  and 
continuity,  chiefly  numerical,  of  the  chromosomes  from  par- 


Inheritance  Materials  of  Germ-Cells  85 

cnt  to  offspring;  the  apparent  equality  (it  should  never  be 
forgotten  that  the  dogma  of  equality  does  not  rest  on  rigor- 
ous quantitative  investigation)  of  the  chromosomes  in  the 
male  and  female  germ-cells;  and  the  assumption  that  in 
tlie  male  germ-cells  the  chromosomes  alone  are  concerned 
in  fertilization.  Kven  if  these  groups  of  assumed  fact  were 
established  with  absolute  certainty  they  would  fall  far  short 
of  being  direct  and  final  proof  that  chromatin  is  tlie  only 
hereditary  substance.  That  this  is  true  must  be  ap})arent 
to  all  well-informed,  carefully  thinking  biologists.  The 
most  important  grounds  for  this  inconclusiveness  are  in- 
volved in  the  facts  and  arguments  set  forth  in  the  preced- 
ing pages,  but  they  may  be  summarized  here,  and  in  addi- 
tion two  other  grounds  may  be  pointed  out. 

First  and  foremost,  in  my  opinion,  is  the  general  trutli 
that  chromosomes  or  even  chromatic  substance  can  not  pos- 
sibly be  recognized  as  the  sole  bearers  of  hereditary  sub- 
stance, because  the  evidence  is  enormous  in  quantity  and 
direct  and  indisputable  in  quality  that  other  substances  })ar- 
ticipate  actively  in  the  production  of  hereditary  attributes. 
There  is  no  way  of  escaping  this  conclusion  except  by  nar- 
rowing the  definition  of  heredity  for  the  very  purpose  of 
bringing  it  within  the  scope  of  the  chromosome  theory  of 
hereditary  substance.  The  scope  and  fundamentalit}^  of 
this  aspect  of  the  problem  is  sufficiently  dwelt  upon,  we 
may  hope,  in  what  has  gone  before. 

The  two  additional  grounds  for  skepticism  as  to  the  con- 
clusiveness of  the  cytological  evidence  will  now  be  pointed 
out.  First,  as  to  the  evidence  from  the  individuality  and 
continuity  of  the  chromosomes.  All  tliat  any  carrful 
thinker  claims  or  can  claim  for  this  evidence  is  tliat  the 
individuality  and  continuity  of  chromosomes  as  observed 
are  what  might  be  expected  if  they  were  the  germinal  i\v- 
pository  of  the  organism's  hereditary  attributes.  Antl  the 
question    is    constantly    and    naturally    asked,    what    other 


86  The  Unity  of  the  Organism 

meaning  can  the  whole  remarkable  series  of  phenomena  of 
maturation  and  fertilization  have  than  that  attributed  to 
them  by  the  chromosome  theory?  My  rejoinder  to  this 
argument  may  begin  with  a  reply  to  the  question.  As  long 
as  knowledge  of  the  chemistry  and  physiology  of  cells — 
germ-cells  with  the  rest — is  as  fragmentary  and  inconclusive 
as  it  now  is,  certainty  as  to  the  meaning  of  the  phenomena 
mentioned  is  out  of  tlie  question.  However,  it  would  seem 
quite  probable  that  they  are  concerned  primarily  with  the 
nutritive  and  assimilative  processes  of  the  cell  and  only 
derivatively  with  heredity.  Furthermore,  the  question  asked 
may  well  be  paired  off  with  another  of'  similar  character, 
namely,  what  is  the  meaning  of  the  almost  if  not  quite  com- 
plete breaking  up  and  disappearance  of  the  chromosomes  in 
the  so-called  resting  stages  of  the  immature  germ-cells,  this 
being  accompanied  by  the  dissolution  of  the  nuclear  mem- 
brane so  as  to  allow  the  freest  possible  commingling  of  the 
whole  nuclear  contents  with  the  cytoplasm  of  the  cell.^^  Have 
we  not  at  least  as  much  factual  right  to  suppose  that  during 
this  mixing  of  substances  the  chromatin,  and  so  later  the 
chromosomes,  are  influenced  by  the  cytoplasm,  as  that  the 
reverse  influence  takes  place  .'^  Is  it  not  entirely  possible 
that  this  process  is  one  of  the  very  means  or  occasions  of 
impressing  the  chromosomes  with  the  attributes  of  the  or- 
ganism which,  as  we  have  seen,  apparently  must  take  place 
whether  acquired  charactei's  are  ever  inherited  or  not.^ 

And  now  as  to  the  argument  from  the  assumed  exclusive 
participation  of  the  chromosomes  of  the  male  germ-cell  in 
fertilization.  First  of  all,  it  can  not  be  admitted  for  a 
moment  that  the  chromosomes  are  proved  to  be  as  ex- 
clusively the  fertilizing  agents  as  they  are  generally  as- 
sumed to  be.  Even  in  such  extreme  cases  of  seeming  ex- 
clusiveness  of  chromosomal  participation  as  that  claimed 
by  Strasburger  for  the  pollen  cells  of  some  plants,  neither 
Strasburger  nor  any  one  else  has  claimed,  so  far  as  I  know, 


Inheritance  Materials  of  Germ-Cells  87 

that  all  other  male  substances  tlian  chromatin  arc  excluded 
as  rigidly  as  would  be  required  by  experiments  in  a  chemical 
hiboratory  designed  to  ascertain  tlie  action  of  a  particular 
chemical  element  or  substance  in  its  purity.  It  is  certain, 
for  example,  that  in  almost  if  not  (juite  all  male  genninal 
elements  in  animals,  a  thin  outer  layer  from  the  cytoplasmic 
part  of  the  spennatid  is  present  on  the  head  of  the  sperma- 
tozoon. Furthermore,  it  is  well  known  that  at  least  the 
"intermediate  piece"  of  the  sperm  tail,  which  is  not  usually 
regarded  as  chromosomal  in  origin,  remains  in  the  Qgg  at 
fertilization.  Nor  is  there  any  good  ground  for  su})posing 
that  the  non-chromatinic  portions  of  the  nucleus  are  ab- 
solutely excluded.  The  almost  certain  presence  in  the  cg^g 
at  fertilization  of  at  least  these  male  substances  other  than 
chromatin  can  by  no  means  be  regarded  as  insignificant  for 
heredity,  especially  if  the  initiator  conception  of  germinal 
material  is  held.  It  seems  to  follow  of  necessity  that  if  the 
fertilizing  substances,  whatever  their  source,  be  conceived 
to  act  in  an  organic  system  of  the  physical  chemistry  sort 
after  the  manner  of  enzymes,  no  such  quantitative  relation 
subsists  between  these  fertilizing  substances  and  the  prod- 
ucts of  organic  growth  as  the  chromosome  theory  implies ; 
nor  can  their  action  be  so  narrowly  localized  in  the  (^gg. 
Their  action  would  be  conceived  to  involve  the  entire  ovum 
ah  initio,  and  not  the  chromosomes  alone. 

Summing  Up  of  the  Findings  Against  the  Chromosome 

Theory 

The  general  result  of  our  critique  is  that  the  whole  at- 
tempt to  interpret  the  physical  basis  of  heredity  in  accord- 
ance with  elementalist  conceptions  has  failed  and  must  con- 
tinue to  fail,  so  far  as  its  main  aim  is  conccrnrd.  We  are 
led  to  see  that  the  germ-plasm  dogma,  no  matter  how  often 
or  how  completely  it  changes  its  nomenclatural  habiliments, 


88  The  Unity  of  the  Organism 

as  in  the  shifting  from  deteYmma7its  to  determiners,  or  from 
determiners  to  ge7is,  or  from  gens  to  factors,  involves  a 
rejection  of  the  conception  that  the  germinal  elements  of 
organisms,  after  being  discharged  are  literally  detached 
parts  of  those  organisms.  This  conception  was  well  on  the 
road  to  incorporation  into  the  great  body  of  established 
biological  truth  when  it  was  headed  ofF  by  Weismann's 
diametrically  opposed  hypothesis  of  germinal  isolation. 

I  would  insist  that  the  defense  of  the  organismal  concep- 
tion in  this  volume  is  really  a  carrying  out  of  such  a  con- 
ception of  the  organism  and  its  germinal  products  as  is 
implied  by  the  old  view  that  the  germ  is  a  part  of  the  parent 
organism.  It  would  hardly  be  possible  to  express  more  sat- 
isfactorily in  a  single  sentence  the  most  inclusive  theo- 
rem, as  it  might  be  called,  the  demonstration  of  which  is  the 
aim  of  the  part  of  this  volume  devoted  to  the  means  by  which 
organisms  propagate  their  kind,  than  the  following  from 
E.  B.  Wilson:  "To  the  modern  student  the  germ  is,  in 
Huxley's  words,  simply  a  detached  living  portion  of  the 
substance  of  a  preexisting  living  body  carrying  with  it  a 
definite  structural  organization  characteristic  of  the 
species."  ^ 

Coupling  this  statement  by  Wilson  with  another  from 
one  of  his  latest  writings,^  to  the  effect  that  we  ought  to 
drop  the  term  determiner  because  in  reality  what  it  means 
is  differential,  I  call  attention  to  the  fact  that  the  "dif- 
ferential factor"  of  the  later  statement  and  the  "definite 
structural  organization  characteristic  of  the  species"  of 
the  earlier  statement  are  in  essence  one  and  the  same.  The 
only  difference  is  that  in  the  earlier  statement  it  is  the 
whole  germ-cell  that  is  recognized  to  be  a  detached  part  of 
the  organism,  while  the  later  statement  can  be  brought 
down  to  the  chromosomes  because  of  the  greater  refinement 
of  knowledge  attained  since  the  earlier  one  was  made.  The 
point  I  wish  to  make  stand  forth  with  the  greatest  possible 


Inheritance  Materials  of  Germ-Cells  89 

boldness  is  tliat  the  germ-cell  chroinosomcs  may  properly 
enough  be  said  to  be  differential,  if  only  one  never  loses 
sight  of  the  fact  that  they  are  differential  in  no  other  sense 
than  are  any  other  particles  or  substances  of  the  germ- 
cells  or  any  other  cells  which  participate  in  the  production 
of  species-attributes. 

Brief  Reference  to  the  Untoxcanl  Implications  of  the  Germ- 
plasm  Conception  of  Heredity 

The  somewhat  laborious  task  of  exhibiting  the  difference 
between  conceiving  the  phj^sical  basis  of  heredity  from  the 
elementalist  and  from  the  organismalist  standpoints  may 
well  be  brought  to  a  close  by  calling  attention  to  the  impli- 
cation of  the  two  conceptions  as  applied  to  heredity  in  man 
himself.  Looked  at  from  this  direction  the  germ-plasm 
dogma  is  seen  to  be  chargable  with  the  grave  offence  of 
having  added  its  weight  to  a  conception  of  human  life,  the 
overcoming  of  which  has  been  consciously  or  unconsciously 
man's  aim  throughout  the  whole  vast  drama  of  his  hard, 
slow  progress  from  lower  to  liigher  levels  of  civilization — 
the  conception  that  his  life  is  the  result  of  forces  against 
which  his  aspirations  and  efforts  are  impotent.  As  ap- 
plied to  man  this  form  of  fatalism  is  no  less  sure  and  no  less 
dire  in  its  tendencies  than  have  been  any  of  the  innumerable 
theistic  forms  of  fatalism  that  have  prevailed  through  the 
centuries.  It  is  almost  certain  that  the  ardor  with  which 
Eugenics  has  been  espoused  by  several  biologists  is  due 
in  considerable  measure  to  the  fact  that  they  have  felt 
more  or  less  definitely  this  sinister  im])lication  of  the  theory, 
and  have  turned  to  Eugenics  as  the  only  weapon  against 
its  evil  forebodings.  The  germ-plasmic  eugenist  virtually 
says,  "Yes,  indeed  is  man  a  reasoning,  willing,  as jn' ring 
animal,  but  all  his  activities  In  these  ways  are  futile  so  far 
as   the   race   as    a   whole   is    concerned,   except    as    they    are 


90  The  Unity  of  the  Organism 

brought  to  bear,  extrinsically  and  operatively  rather  than 
organically,  on  the  Germ-Plasm."  This  form  of  the  Eu- 
genic idea  corresponds  in  spirit  to  the  propitiative  offerings 
of  primitive  religion.  It  aims  to  mollify  by  human  agency 
powers  that  act  upon  men's  Hves,  but  which  are  in  them- 
selves largely  extraneous,  largely  evil,  and  wholly  irre- 
sponsible. 

What  eugenists  of  this  school  have  failed  to  see,  evidently, 
is  that  even  were  unit-factors  as  differentiate  from  one 
another  in  heredity  as  the  extremest  Mendelist  conceives 
them  to  be,  and  that  even  were  the  germ-plasm  improved  up 
to  the  level  of  his  highest  hopes,  his  results  in  terms  of  ac- 
tual human  lives  and  social  conditions  would  be  distressingly 
meager.  They  would  be  so,  because  whether  unit-factors 
exist  independently  in  heredity  or  not,  they  certainly  do 
not  exist  thus  independently  in  development  and  function. 
In  these  ways  they  interact  upon  one  another  in  the  most 
vital  manner,  as  physiology,  especially  of  the  internal  se- 
cretions and  the  nervous  system,  and  as  physiological  and 
social  psychology  are  rapidly  and  conclusively  demon- 
strating. 

We  thus  end  our  examination  of  the  means  by  which  or- 
ganisms produce  others  of  their  kind  with  the  conclusion 
that  the  material  through  which  reproduction  is  accom- 
phshed  is  in  the  most  vital  way  part  and  parcel  of  the 
organism,  that  is,  that  the  germ-cells  are  somehow  stamped 
through  and  through,  potentially,  with  the  characteristics 
of  the  kind,  or  race,  or  species  to  which  the  producing  or- 
ganism belongs.  And  with  this  we  are  ready  to  pass  to  the 
examination  of  those  integrative  phenomena  of  the  organ- 
ism generally,  one  manifestation  of  which  is  this  very  nature 
of  the  germ-cells. 

REFERENCE  INDEX 

1.  Marshall    292         4.  Riddle     10 

2.  Hertwig,  R.   ('12)    75         5.  Wilson,  E.  B.    ('14) 351 

3.  King    232        6.  Wilson,  E.  B.   ('00) 7 


PART   II 

THE    CONSTRUCTIVE    SIDE    OF   THE    ORGANISMAL 

CONCEPTION 


Chapter  XV II 
GROWTH   INTEGRATION 

The  Field   to   he  Covered  hy   the  Constructive  Discussion 

ACCEPTING  the  inevitable  destructive  result  of  our  cri- 
tique of  the  clemcntalist  standpoint,  that  the  attempt 
to  interpret  living  beings  in  the  terms  of  their  constituent 
parts  alone  always  leads  to  partial  failure  and  disappoint- 
ment, or  to  the  worse  result  of  illusionment  as  to  the  trust- 
worthiness of  the  explanations  proposed ;  and  accepting  the 
constructive  result  that  everything  in  the  critical  study  tends 
to  show  that  no  part  of  any  organism  can  be  rightly  inter- 
preted except  as  part  of  an  individual  organism,  this  indi- 
vidual being  in  turn  interpreted  as  a  member  of  a  taxonomic 
group,  it  is  revealed  that  we  are  only  on  the  tlireshold  of  tlie 
positive,  the  constructive  side  of  our  general  enterprise.  Even 
though  the  conclusion  be  unescapable  that  the  living  organ- 
ism someliow  acts  causally  on  its  parts,  the  problem  still 
remains  as  to  the  modus  operandi  of  tliat  acting.  The  "some- 
how" which  came  to  us  as  an  incident  of  our  critical  study 
has  yet  to  be  inquired  into. 

Stated  more  specifically  the  task  now  Inforc  us  is  Hiat  of 
examining  closely  and  systematically  the  interdependences 
among  the  parts  of  the  individual  organism.  Although  these 
interdependences  are  among  the  most  obvious  and  general 
of  all  organic  phenomena  such  an  examination  of  them  biol- 
ogy has  not  yet  made  systematically.  Indeed — and  here  is 
one  of  the  most  vital  things  for  us  to  see — a  cardinal  charge 
against  the  elementalist  standpoint  is  that  in  its  very  nature 

93 


94  The  Unity  of  the  Organism 

it  not  only  does  not  encourage,  it  actually  stands  against 
such  examination.  Its  opposition  to  comprehensiveness  and 
systematization  is  profound  and  essential. 

Our  examination  will  begin  with  a  single  brief,  two-parted 
definition :  The  structural  and  functional  interdependence 
found  to  exist  among  the  parts  of  an  organism  we  call  hio- 
integratedness;  and  the  process  of  moving  on  from  grade 
to  grade  of  interdependence  among  the  differentiating  parts 
which  constitutes  ontogenesis  in  the  individual  we  call  hio- 
integration. 

Four  Types  of  Bio-integration  to  Be  Treated 

In  the  present  state  of  knowledge  and  for  the  discussion 
now  before  us  four  types  or  kinds  of  bio-integratedness  and 
bio-integration  may  be  recognized  as  pertaining  to  the  in- 
dividual organism : 

1.  Growth  integration,  most  obvious  in  graded  meristic 
series,  but  also  expressed  in  the  "axial  gradients"  of  Child. 

2.  Chemico- functional  integration,  known  so  far  chiefly 
in  connection  with  internal  secretions. 

3.  Neural  integration,  comprising  the  interdependences 
among  the  parts  of  the  nervous  system,  and  the  involvement 
with  this  of  the  muscular,  glandular  and  other  organs. 

4.  Psychic  integration,  very  closely  connected  with  neural 
integration,  but  approached  from  the  side  of  the  totality  of 
activities  of  living  beings  rather  than  from  the  side  of  nerve- 
organ  activity,  and  so  taking  cognizance  of  a  vast  number 
of  phenomena  not  yet  definitely  cor relat able  with  neural 
phenomena. 

The  full  presentation  of  facts  and  arguments  under  these 
four  heads  would  reach  far  beyond  the  limits  set  for  the 
present  work.  We  are,  consequently,  obliged  to  restrict 
ourselves  to  a  small  portion  of  the  best  established  and  most 
compelling  evidence  under  each  head. 


Growth  Integration  95 


Graded  Repetitive  Series  as  Integrative  PJienojrwna 

This,  perhaps  the  simplest  form  of  integrational  phenom- 
ena known  to  biology,  is  seen  almost  everywhere,  but  shows 
itself  most  typically  and  strikingly  in  plants  and  in  many 
lower  animals.  Reference  is  made  to  the  gradation  in  the 
repetitive  or  meristic  parts  appearing  in  so  many  organisms. 
The  most  obvious  criterion  of  sucli  gradation  is  the  rela- 
tive size  of  the  parts,  but,  as  we  shall  see  later,  there  is  con- 
siderable reason  for  supposing  the  gradation  is  not  re- 
stricted to  size.  The  few  examples  to  which  space  can  be 
given  are  selected  to  represent  as  wide  a  range  as  possible 
of  the  phenomena  under  consideration. 


Illustrations  from  Ani/mals 

\         The  lancelets,  fish-like  animals  of  the  genus  Amphioxiis, 
may  be  noticed  first  (Figure  55).     It  will  be  obsei-^ed  that 


"^^        <*»*a  ^- 


FIGURE    55 — SIDE    VIEW    Or    AMPIIIOXUS     ( AFTER    PARKKR    4c    IIASWELl).       Iicll., 

notocliord.    cir.,  cirri,    or.hd.,  oral  hood,    myoni.,  myomeres,    dors.fr., 
dorsal  fin  rays,     cd.f.,  caudal  fin.     gon.,  gonads 

the  creature  tapers  oft*  toward  botli  ends  and  that  the  series 
of  metameres,  myom,  usually  called  myomeres  because  they 
compose  the  main  body-musculature,  diminish  not  only  in 
dorso-ventral  measurement  from  near  the  middle  each  way, 
but  are  also  thickest  in  the  mid-region  and  become  thinner 
as  they  progress  toward  each  end. 

Something  of  this  size  scheme  of  body-parts  is  very  com- 


96 


The  Unity  of  the  Organism 


mon  in  the  animal  kingdom.  Figure  5Q,  a  photograph  of 
the  skeleton  of  a  python  (see  frontispiece),  shows  in  a  gen- 
eral way  the  size-relations  of  metameric  skeletal  parts  in  a 
higher  vertebrate.  Something  of  the  extent  to  which  the 
proportionality  of  parts  of  the  individual  metameres  is  car- 
ried out  in  this  skeleton  is  shown  by  tabulating  a  series  of 
measurements  of  the  parts : 


Position  of 

vertebra 

L.V. 

T.V.       T. 

ex.zyg. 

H.D.S. 

W.D.S. 

L.R. 

5 

6  mm 

5  mm. 

11   mm.   15  mm.     4  mm. 

25  mm. 

50 

11 

16 

23 

11 

8 

48 

100 

13 

17.0 

28.5 

9 

7 

79 

150 

13 

16.5 

26.5 

7.5 

7 

82.5 

200 

11.5 

15 

22.5 

5.5 

7.5 

Q9.5 

250 

10 

10.5 

16.5 

5 

5.5 

52.5 

300 

6 

5 

7 

4 

2 

0 

327 

4 

3 

4 

2 

2 

0 

"Position  of  vertebra"  refers  to  the  serial  number,  beginning 
with  the  head-end,  of  the  vertebra  measured.  Legend:  L.V., 
Length  of  vertebra,  measured  from  the  posterior  edge  of  one 
dorsal  spine  to  the  anterior  edge  of  the  one  next  behind  it.  T.V., 
thickness  of  vertebral  centrum  in  its  thinnest  part,  i.e.,  near  the 
middle.  T.  ex.  zyg.,  thickness  of  vertebra  at  extreme  of  posterior 
zj^gopophyses  (articulating  processes).  H.D.S.,  height  of  dorsal 
spine.     W.D.S. ,  width  of  dorsal  spine.     L.R.,  length  of  rib. 


The  starfishes  are  another  class  of  animals  which  exhibit 
beautifully  this  size  gradation  of  repeated  parts,  both  their 
"tube-feet"  and  the  calcareous  skeletal  supports  being 
graded  proportionately  to  the  tapering  arms  of  the  animal. 
The  following  table  presents  a  series  of  measurements  of  the 
two  organ  systems  just  mentioned,  from  a  single  arm  of 
Astrospecten  calif ornicus.  The  dimensions  are  in  millimet- 
ers, and  the  series  proceed  from  the  proximal  to  the  distal 
end  of  the  arm. 


Growth  Integration  97 

TABLE 

Series  Tube-feet       Ambulacral  plates  Adambulacral  plates 

number    Length  Length   Width  Thickness  Length   Width    Thickness 

5  3.9  6.4  3.1  1.2  3.8  S.Q  1.0 

10  3.7  3.9  2.7  1.3  S.5  2.4  1.2 

15  S.5  3.5  2.4  1.3  2.8  2.2  1.4 

20  3.1  2.6  2.0  1.2  2.4  2.0  1.2 

25  2.2  2.3  2.0  1.0  2.1  1.9  1.1 

30  2.1  2.1  1.9  0.9  1.9  1.7  1.2 

35  2.3  1.8  1.6  1.0  1.7  1.3  1.1 

40  1.7  1.2  1.1  0.9  1.4  1.0  1.0 

45  1.8  0.8  1.0  0.6  1.0  0.8  0.9 

50  1.3  0.7  0.7  0.5  0.7  0.7  0.6 

55  0.9 

60  0.8 

65  0.7 

68  0.5 

Not  only  do  these  graded  meristic  series  appear  in  the 
individual  makeup  of  a  great  range  of  animal  species,  hut 
they  occur  in  the  colonies  of  many  species  in  which  aggrega- 
tions are  produced  by  budding.  Sometimes,  as  in  many  al- 
cyonaria,  the  size  gradations  are  very  obvious,  wliile  in  otlier 
groups  the  distinctions  are  so  small  as  to  be  discoverable 
only  by  close  quantitative  study.  An  example  of  this  latter  is 
furnished  by  the  plumularian  hydroids.  A  typical  colony  of 
the  genus  here  studied,  Torrey  writes,  "closely  resembles  a 
feather,  of  which  the  shaft  is  represented  by  the  stem  and  the 
veins  by  the  two  ranks  of  alternating  branchlets,  or  hydro- 
cladia,  corresponding  to  barbs.  Each  hydrocladiuni  is 
divided  by  more  or  less  definite  nodes  into  internodes  and 
bears  on  one  aspect — the  same  in  all  hydrocladia — a  compact 
series  of  hydranths,  one  to  each  internode."  ^ 

Without  entering  into  the  tabular  and  graphic  details 
contained  in  this  study,  the  author's  summarized  statement 
concerning  one  of  the  tables  will  suffice:  "It  will  be  seen  from 
the  table  that,  as  the  tip  of  the  colony  is  approached,  not 


98 


The  Unity  of  the  Organism 


only  do  the  hydrocladia  possess  fewer  and  fewer  hydroth- 
ecae,  but  the  dimensions  of  the  latter  through  the  mesial 
nematophore  reaches  its  minimum  more  and  more  rapidly. 
Since  the  hj^drothecae,  once  formed,  do  not  enlarge  with  age, 
it  is  clear  that  for  such  colonies  as  this,  there  is  a  limit  of 
growth  and  a  specific  form."  ^ 

This    correlation    and   proportionality    among    repetitive 
parts  is  frequently  observed  within  the  bounds  of  particular 


S^ 


-V— V 


a. 

figure    57 — tentacle    of    halocynthia    johnsoni    (schematic;    after 
ritter). 
a,  axis,    a',  axis  of  branch,    b',  b",  primary  and  secondary  branches. 


organs,  as  for  example,  in  the  branching  tentacles  occurring 
in  various  groups.  A  detailed  study  of  one  of  these  cases 
was  made  by  me  some  years  ago  on  the  tentacles  of  an  as- 
cidian.  Figure  (pi.  12,  figure  13)  of  this  study,  sup- 
plemented by  the  following  statement,  illustrates  the 
point.  "Although  this  figure  is  diagrammatic  in  a  way,  it  is 
accurate  as  to  numbers  of  branches.  The  positions,  too,  of 
all  the  branches  and  length  of  the  primaries  were  deter- 
mined by  micrometer  measurements,  and  the  secondaries  were 
drawn  as  accurately  as  possible."  ^  For  the  rest,  the  figure 
(Figure  57)  tells  its  own  story. 


Growth  Integration 


99 


Illustrations  from  Plants 

But  it  is  in  tlie  plant  world  that  those  graded  scries  of 
homonynious  parts  in  individual  organisms  are  most  strik- 
ingly seen.  It  occurs  in  what  is  perha])s  its  most  typical, 
least  modified  expression  in  the  arrangement  and  size  rela- 
tion of  parts  in  the  leaves  of  many  ferns  and  palms.  Hut 
the  compound  leaves  of  innumerable  flowering  ])lants  illus- 
trate it  very  beautifully.  Figures  58,  59,  and  60  {.icacia, 
Vicia  and  Cassia)  show  three  types  of  compound  leaves 
based  on  the  mode  of  gradation  of  the  leaflets.     These  might 


FIOURE    58.       ACACIA    ELATA. 


100 


The  Unity  of  the  Organism 


be  described  as  the  bi-gradient,  the  direct  gradient  and  the 
reverse  gradient  types,  depending  on  whether  the  gradation 
is  from  the  mid-region  of  the  axis  both  ways  (figure  58),  from 
the  proximal  toward  the  distal  end  (figure  59),  or  from  the 
distal  toward  the  proximal  end  (figure  60). 


FIGURE    59.      VICIA    GIGANTEA. 


FIGURE    60.       CASSIA    SP. 


Almost  all  simple  leaves  of  seed  plants  show  something  of 
the  same  scheme.  As  examples,  typical  elliptical-entire 
leaves  of  the  elm  and  poplar  and  such  typical  lobed  leaves  as 
those  of  most  oaks  may  be  pointed  to. 

Nearly  every  twig  of  a  tree  which  represents  a  single 
annual  growth  impulse,  in  cases  where  the  growing  period  is 
restricted  to  a  small  part  of  each  year,  presents  a  size  gra- 
dient in  the  leaves  distributed  along  the  axis.  A  particularly 
striking  illustration  of  this  is  furnished  by  the  California 


Growth  Integration 


101 


coast  redwood,  Sequoia  sernperiirens  (figure  f)l),  wliere  the 
new  segment  is  sliort,  is  added  end  on  to  the  one  before  it 
until  a  considerable  succession  of  segments  is  ])ro(liifed,  and 
where  the  leaves  are  retained  for  several  years.  That  each 
segment  in  these  leaves  is  an  annual  production  is  not  cer- 
tain, probably  several  segments  being  sometimes  formed  in  a 


FIGURE    61.      SEQUOIA    SEMPERVIHENS. 

single  season;  but  however  that  may  be,  that  growth  occurs 
in  a  series  of  impulses,  each  of  which  is  sharply  recorded 
in  the  size  gradations  of  the  repeated  parts,  is  obvious 
enough. 

It  is  a  familiar  fact,  too,  that  in  many  plants  a  similar 
quantitative  gradation  of  the  reproductive  parts  along  an 
axis  occurs,  but  the  extent  to  which  this  scheme  pervades 
the  constituents  of  the  members  of  the  series,  even  to  tlie 
seeds,  appears  not  to  have  attracted  much  interest  on  the 
part  of  botanists.  To  illustrate  this  point  I  present  a  single 
set  of  measurements,  one  of  many   which   I   have  collected, 


102  The  Unity  of  the  Organism 

of  parts  of  the  fructiferous  organs  of  plants.  These  meas- 
urements are  of  Frasera  perryi,  an  abundant  annual  in 
southern  California  and  rather  specially  favorable  for  such 
a  study  in  that  the  fruit  stalk  is  single  in  each  plant,  stands 
up  intact  and  rigid  after  it  is  fully  ripe  and  dry,  and  is  al- 
most mathematically  regular  in  the  disposition  of  its  parts. 
The  table  was  compiled  from  measurements  of  a  single  plant, 
and  three  measurements  pertaining  to  each  seed  vessel  are 
given,  namely,  the  length  of  the  interval  on  the  main  axis 
between  each  two  vessels,  the  length  of  the  pedicels  which 
bear  the  vessels,  and  the  length  of  the  vessels  themselves. 
The  measurements  are  all  in  millimeters.  Several  other  di- 
mensions might  have  been  taken,  which  would  almost  cer- 
tainly have  produced  similar  results. 


Series 

Length  of 

Length  of 

Length  of 

slumber 

Internode 

Pedicel 

Capsule 

1 

27 

36 

21 

2 

19 

29 

18 

3 

19 

32 

18 

4 

22 

34 

16 

5 

23 

29 

16 

6 

19.5 

28 

17 

7 

18.5 

25 

17.5 

8 

17 

25.5 

17.5 

9 

19 

25 

18 

10 

18 

25 

16 

11 

18 

21 

16.5 

12 

15 

21 

16 

\S 

15 

22 

13.5 

14 

15 

22 

14 

15  15  21.5  13 

That  these  gradations  hold,  at  least  in  some  plants,  even 
to  the  seeds  is  certain  as  tlie  following  tabulation  of  the 
weight  of  seeds  from  different  parts  of  the  seeding  axis  of  a 
wild  mustard  plant  (Brassica  nigra)  shows.  The  figures 
were  compiled  from  the  weights  of  seeds  taken  from  groups 


arc  in  grains. 

Total  weight 

Base 

.903 

Middle 

.694> 

Tip 

.330 

Grorcth   Infcgration  103 

of  twenty  ca])sule.s  from  the  bases,  niiddK-  |)orHons,  and  dis- 
tal ends,  respectively,  of  six  such  stalks.     The  weights  given 

Number  of  Seeds     Av.  Wt.  per  Seed 

950  .0009.0 

733  .00088 

525  .000()3 

This  mere  glance  at  an  exceedingly  common  phenomenon 
in  living  nature  must  suffice  for  the  present. 


Justification  for  Bringing  All  These  Phenomena  Under  One 

Head 

Probably  about  the  first  question  that  most  persons  would 
raise  concerning  what  we  have  presented  would  be  as  to  how 
far  the  series  dealt  with  have  anything  to  do  with  one  an- 
other. Especially,  we  may  apprehend,  would  most  biologists 
question  the  justifiability  of  bringing  together  the  meristic 
phenomena  in  animals  and  the  repetition  of  parts  in  plants. 
If  such  a  collocation  of  phenomena  must  be  justified  on  the 
basis  of  known  causal  factors,  then  undoubtedly  is  justifica- 
tion impossible  in  the  present  state  of  knowledge.  But  justi- 
fication of  this  sort  is  not  called  for  by  the  point  now  oc- 
cupying us.  What  concerns  us  at  present  is  the  quite  for- 
mal fact  that  when  any  lot  of  homonymous  objects  fall  into 
a  quantitatively  graded  series  the  members  of  that  series 
liave  a  fixed  relation  to  the  series  as  a  whole.  They  n\\-  not 
interchangeable  with  one  another.  Kach  is  a  function, 
mathematically  speaking,  of  its  set  or  series.  Vertibra  /;/ 
of  the  python's  skeleton,  myotome  ni  of  the  amphloxus  Ixxly, 
tube-foot  7n  of  the  starfish  arm,  branchlet  in  of  the  ascidian 
tentacle,  leaflet  m  of  the  vetch  leaf  or  of  the  redwood  shoot, 
seed-vessel  m  of  Frasera,  seed-lot  m  of  the  mustard  plant, 
and   m,   or   any   other  member  you  choose   from   any   other 


104  The  Unity  of  the  Organism 

series  whatsoever,  is  a  determinate  thing;  it  is  what  it  is 
partly  because  of  its  position  in  the  series  regardless  of 
whether  the  physical  or  other  producing  agent  of  the  dif- 
ferent series  be  the   same  or  wholly   different. 

Even  this  purely  structural  formal  basis  establishes 
the  fact  of  a  measure  of  integratedness  for  all  individual 
organisms  in  which  the  phenomenon  appears.  But  to  leave 
the  subject  at  that  would  be  superficial  and  unsatisfactory 
indeed.  However,  reflection  makes  it  almost  certain  that 
there  is  some  sort  of  causal  basis  for  the  phenomena.  This 
conclusion  follows  first  from  the  fact  that  the  series  result 
from  the  growth  of  the  organism,  and  second  from  the  cer- 
tainty, at  least  in  many  cases,  that  the  continuance  of  life 
of  the  individual  involves  the  maintenance  of  the  series,  this 
in  turn  involving  some  measure  of  metabolic  interdepen- 
dence among  the  members  of  the  series. 

Attempted  Causal  Explanation  of  These  Series 

For  establishing  the  general  truth  of  this  type  of  inte- 
gration we  need  not,  in  strictness,  go  any  further  than  we 
have  gone.  Nevertheless,  the  importance  of  the  subject 
justifies  a  few  remarks  on  attempts  that  have  been  made  to 
explain  the  series  causally.  The  best  known  of  these  comes 
from  botanists,  and  conceives  that  the  diminishing  series  of 
leaves  and  other  structures,  seen  with  more  or  less  distinct- 
ness almost  universally  among  plants,  is  due  to  the  increas- 
ing remoteness  of  the  successive  parts  from  the  roots  of  the 
plant,  that  is,  from  the  main  source  of  the  plant's  food.  It 
is  obvious,  however,  that  this  explanation  is  not  of  general 
application,  since  in  animals  the  food  does  not  come  from  a 
root  system  which  anchors  the  organism  to  its  food-yielding 
medium.  Nor  is  it  possible  to  bring  the  series  in  all  animals 
into  correlation  with  a  blood  circulatory  system,  as  their 
existence  in  many  coelenterates,  hydroids  and  alcyonarians 


Growth  Integration  105 

for  cxanipk',  wlicre  mo  circulation  exists,  sliows.  It  is  al- 
most certain,  too,  that  the  series  occur  in  iiiaiiv  plants  that 
liave  no  sap  system  such  as  is  assumed  by  the  ])hvsiolo^ical 
explanation  above  indicated.  Many  of  the  marine  al^ae 
come  under  this  head,  a  striking  example  of  which  is  the  kel() 
M<icrocystis  pyrifera  of  the  western  coast  of  hotli  Americas. 
'J1iat  the  laminje  of  this  plant  fall  into  a  beautiful  direct 
gradient  series  is  a  fact  whicli  can  not  escape  the  notice 
of  any  one  who  sees  them.  The  question  of  whether  each 
streamer  of  laminae  reaches  finally  and  necessarily  a  limit  of 
growth  in  which  the  size  scries  is  present  is  not  so  certain, 
but  from  considerable  attention  to  the  question  I  am  almost 
sure  this  is  the  case,  although  the  point  needs  more  study. 
Another  interesting  and  probably  useful  course  of  reason- 
ing about  organic  growth  attempts  to  connect  tlie  results  of 
growth  with  autocatalytic  chemical  action.  Although  these 
attempts  have  not,  so  far  as  I  am  aware,  taken  special  cog- 
nizance of  the  natural  size  series  which  are  occu})ying  us, 
but  have  been  concerned  with  the  weights  or  volumes  of  or- 
ganisms at  various  stages  of  growth,  there  is  little  doubt 
that  the  phenomena  we  have  been  considering  are  closely 
connected  with  those  dealt  with  in  these  attempts.  T.  Brails- 
ford  Robertson  seems  to  have  devoted  more  thought  to  this 
matter  than  any  one  else.  The  following,  taken  from  the 
summary  of  conclusions  found  in  his  original  ])a})er,  ])re- 
sents  the  most  essential  parts  of  his  theory:  "(1)  In  any 
particular  cycle  of  growth  of  an  organism  or  of  a  particular 
tissue  or  organ  of  an  organism  the  maximum  increase  in  vol- 
ume or  in  weight  in  a  unit  of  time  occurs  when  the  total 
growth  due  to  the  cycle  is  half  completed.     (  ^i  )  Any  pai'tieu- 

X 

lar  cycle  of  growth  obeys  the  fornuila  lay -~  K(t-t,), 

A.  X 

where   x  is    the   amount    (in    weight    or   volume)    of   growth 

which  has  been  attained  in  time  f,  A  is  the  total  amount  of 

growth  attained  during  the  cycle.  A'  is  a  constant,  and  ^  is 


106  The  Unity  of  the  Organism 

the  time  at  which  growth  is  half  completed."  ^ 

Assuming  that  the  growth  of  an  amphioxus,  let  us  say,  to 
adulthood  represents  a  growth  cycle  of  this  statement^  that 
the  production  of  somites  begins  with  the  most  anterior  one 
and  proceeds  toward  the  tail,  and  that  the  successive  growth- 
increments  (corresponding  to  oc  in  tlie  formula)  are  regis- 
tered in  the  somites  as  we  find  tliem,  then  the  animal's  bodv 
as  exhibited  by  its  myomeres  would  correspond  fairly  well 
to  Robertson's  statement  under  (1),  as  would  several  of  the 
other  growth  series  we  have  glanced  at,  and  as  would  also 
great  numbers  of  series  presented  by  ordinary  plants  and 
animals. 

The  formula  for  growth  contained  in  (2)  is,  according 
to  Robertson,  "such  as  would  be  expected  to  hold  good  were 
growth  the  expression  of  an  autocatalytic  chemical  reac- 
tion." Assuming  the  general  correctness  of  these  state- 
ments, no  one  interested  in  the  larger  problems  of  organic 
growtli  could  hesitate  to  believe  that  they  must  be  impor- 
tant in  some  way. 

However,  that  the  relations  shown  do  not  prove  that  au- 
tocatalytic chemical  activity  is  a  cause  of  growth  in  any- 
thing more  than  a  subordinate,  contributory  way,  is  obvious^ 
on  reflection.  In  the  first  place,  Robertson  himself  has 
pointed  out,  in  substance,  that  such  action  says  nothing 
about  the  particular  shape  which  the  mass  of  transformed 
substance  takes,  but  since  some  characteristic  configuration 
or  shape  is  fundamental  to  all  organic  growth,  the  entities 
for  which  A  and  x  stand  in  the  formula  are  really  only  ab- 
stractions. Altliough  the  formula  may  apply  approximately 
to  a  great  many  organisms,  it  will  apply  to  none  exactly, 
except  by  cliance  to  a  very  occasional  one.  This  is  the  gen- 
eral form  of  criticism,  illustrations  of  which  are  seen  in  the 
fact  that  in  the  series  of  direct  and  inverse  gradients  shown 
in  the  vetch  (figure  59)  and  Cassia  (figure  60),  respectively, 
the  formula  appears  not  to  apply  at  all.     The  general  pur- 


Growth  Integration  107 

port  of  tlie  strictures  here  placed  upon  tlio  value  of  this 
explanation  of  growth  is  well  l)r()u<;i»t  out  hy  Mocser,  who 
says,  probahly  with  literal  trutli fulness,  "'One  will  not  find 
two  germinating  plants  {Keimp'jlanzen)  which  would  have 
exactly  the  same  growth  curve,  even  though  they  proceed 
from  seed  of  absolutely  tlie  same  weight  and  grow  under 
exactly  the  came  conditions."  ^ 

The  kernel  of  this  criticism  is  that  even  tliough  it  should 
be  established,  as  very  likely  it  will  l)e,  that  autocatalytic 
action  is  an  essential  factor  in  all  growth,  it  can  be  a  causal 
explanation  in  only  a  partial  and  subordinate  sense.  This  is 
so  because  although  TT  is  a  constant  for  a  particular  indi- 
vidual as  observed,  it  assumes  different  values  for  different 
groups,  partly  at  least  because  no  account  is  taken  of  size 
or  configuration.  Moreover,  even  if  these  factors  were  con- 
sidered, there  would  be  nothing  corresponding  to  a  physical 
constant  (depending  only  on  autocatalytic  action),  since 
one  of  the  most  distinctive  things  about  organic  growth  is 
that  it  is  differential,  the  differentials  corres]iondIng  to  the 
taxonomic  group  to  which  the  individuals  belong. 

Even  though  we  are  still  much  in  the  dark  as  to  what  the 
causal  nexus  is  between  the  growth  processes  and  tlie  (|uan- 
titatively  graded  series  so  widely  seen  in  nature,  it  seems 
certain  that  some  such  nexus  exists,  and  that  its  operation 
constitutes  a  true  integ-rational  factor  in  the  individual.  It 
appears,  too,  that  this  type  of  integration  is  about  the 
simplest  and  that  it  accom])anies  tlie  sim])lest  type  of  dif- 
ferentiation, the  two  together  constituting  the  sini])lest  type 
of  organization  above  the  organization  of  the  cell.  W\\\  into 
this  interesting  subject  we  can  not  now  go. 

Axial  Metabolic   (irad'icnts   as    Infrgrathr    Phcnoimua 

For  the  most  thorough  and  sustained  experinieiif al  inves- 
tigation  of   the    2>rimary    integrative    processes    in    growing 


108  The  Unity  of  the  Organism 

organisms  that  has  been  made,  biology  is  indebted  to  C.  M. 
Child.  In  two  recent  volumes  he  has  summed  up  and  sys- 
tematized the  elaborate  researches  prosecuted  by  him  in  this 
field  almost  exclusively  for  fifteen  years,  and  has  presented 
his  conclusions  more  fully  than  in  any  of  his  previous  writ- 
ings. 

The  limitation  set  for  the  present  to  the  constructive  part 
of  our  enterprise  makes  it  impossible  to  do  more  than  touch 
at  a  few  points  the  great  mass  of  experimental  evidence  on 
which  Doctor  Child  bases  his  conclusions.  Fortunatelv, 
however,  the  kernel  of  the  conclusions  can  be  stated  rather 
clearly  in  a  small  space. 

Although  (surprisingly,  it  seems  to  me)  Child  refers 
hardly  at  all  to  the  graded  meristic  series  occurring  in  na- 
ture, to  which  the  preceding  pages  have  been  devoted,  it 
can  hardly  be  doubted  that  the  phenomena  with  which  he 
deals,  and  calls  "axial  gradients,"  come  under  the  same  head 
as  do  those  which  have  been  occupying  us.  The  phenomena 
which  in  the  first  instance  Child  has  been  concerned  with, 
have  been  brought  to  light  mainly  through  studies  on  re- 
generation in  many  lower  animals.  But  the  general  con- 
clusions reached  are  far  broader  than  this ;  indeed  they  ex- 
tend to  well-nigh  the  whole  scope  of  organic  growth,  but 
especiall}'-  to  growth  which  involves  elongation  either  of  the 
whole  organism  or  of  certain  parts  of  organisms.  Thus  the 
head-tail  type  of  individual,  whether  the  body  be  segmented 
as  in  arthropods  and  in  many  worms,  or  unsegmented  as  in 
other  worms  and  in  molluscs,  is  perhaps  the  most  striking 
exemplification  of  the  axial  gradations  with  which  Child 
deals.  The  following  quotation  shows  the  generality  with 
which  he  views  the  matter  from  the  ontogenic  side:  "Gra- 
dients in  rate  of  cell  division,  size  of  cells,  condition  or 
amount  of  protoplasm  in  the  cells,  rate  of  growth,  and  rate 
and  seqvience  of  differentiation  are  very  characteristic  fea- 
tures  of  both   animal   and  plant   development.      Such   gra- 


Growth  Integration  109 

dients  are  definitely  related  to  the  axes  of  the  individual  or 
its  parts,  and  are  evidently  expressions  of  axial  metabolic 
gradients.  While  the  existence  of  such  gradients  indicates 
tlie  existence  of  gradients  in  activity  of  sonic  sort,  the 
various  kinds  of  gradients  are  not  all  necessarilv  pres- 
ent where  metabolic  gradients  exist.  In  some  cases  the  vis- 
ible gradient  may  be  a  gradient  in  rate  of  growth  or  In 
protoplasmic  constitution;  in  still  others  a  gradient  in 
sequence  of  differentiation,  etc.,  and  sometimes  mctaholic 
gradients  exist  without  any  structural  indications  of  their 
presence.  At  best  these  various  kinds  of  gradients  arc 
merely  general  indications  of  differences  in  metabolic  rate 
and  undoubtedly  in  many  cases  the  visible  differences  along 
an  axis  represent  something  more  than  differences  in  meta- 
bolic rate.  The  important  point  is  that  visible  indications 
of  graded  differences  in  metabolic  rate  occur  so  generally 
in  definite  relations  to  the  chief  axes  of  the  body.'' 

One  phase  of  this  general  statement  is  the  developmental 
correlation  that  various  regions  of  the  bodv  in  manv  lower 
animals  have  with  the  head  or  anterior  end,  these  regions 
being  developmentally  dominated,  in  Child's  expression,  by 
the  anterior  end  proportionally  to  the  distance  of  the  re- 
gion from  the  end. 

A  typical  case  is  furnished  by  fiat-worms  of  the  genus 
Planaria,  animals  especially  favorable  for  e\[)eriments  in 
regeneration,  since  they  are  very  hardy  to  laboratory  con- 
ditions and  have  great  powers  of  reconstituting  themselves 
from  pieces  of  various  sizes,  sha])es  and  ])ositions  cut  from 
them.  "Any  piece  of  the  ])lanarian  l)ody,"  says  Child,  '"is 
capable  of  giving  rise  to  all  parts  posterior  to  its  own  level, 
whether  a  head  is  present  or  not,  but  no  piece  is  caj)ahle  of 
producing  any  part  characteristic  of  more  anterior  levels 
than  itself,  unless  a  head  begins  to  form  first."  ^ 

From  a  great  mass  of  experimental  evidence  produced  bv 
Child  and  others  we  have  the  following:  "These  facts  force 


110  The  Unity  of  the  Organism 

us  to  the  conclusion  that  in  such  experimental  reproductions 
there  is  a  relation  of  dominance  and  subordination  of  parts. 
The  apical  or  head  region  develops  independently  of  other 
parts  but  controls  or  dominates  their  development,  and  in 
general  any  level  of  the  body  dominates  more  posterior  or 
basal  levels  and  is  dominated  by  more  anterior  or  apical 
levels."  '^ 

A  really  unique  merit  in  Child's  work  is  the  fact  that  he 
has  given  special  attention  to  the  connection  of  these  axial 
gradients  manifesting  themselves  in  various  structural  and 
functional  ways,  with  the  fundamental  metabolism  of  the 
organism.  Several  methods  of  experimenting  have  been  em- 
ployed to  this  end,  the  one  most  frequently  used  being  what 
he  calls  the  susceptibility  or  survival-time  method.  The  es- 
sence of  this  depends  upon  the  fact,  determined  by  many  ob- 
servers, "that  a  relation  exists  between  the  general  meta- 
bolic condition  of  organisms,  or  their  parts,  and  their  sus- 
ceptibility to  a  very  large  number  of  substances  which  act 
as  poisons,  i.e.,  which  in  one  way  or  another  make  meta- 
bolism impossible,  and  that  difference  in  susceptibility  may 
be  used  with  certain  precautions  and  within  certain  limits  as 
a  means  of  distinguishing  differences  in  metabolic  condition, 
and  more  specifically,   differences   in   metabolic   rate."  ^ 

The  demonstration  of  metabolic  gradients  by  this  method 
depends  upon  the  fact  that  "death  and  disintegration  of  dif- 
ferent parts  of  the  body  usually-  follow  a  regular  sequence," 
this  making  it  possible  "to  determine  the  time,  not  merely  of 
disintegration  of  the  whole  animal,  but  of  the  various  re- 
gions of  the  body."  ^ 

Another  way  of  showing  difference  in  rate  of  metabolism 
in  different  parts  of  the  organism  is  by  the  use  of  the  biom- 
eter,  an  apparatus  for  estimating  minute  quantities  of  carbon 
dioxide,  recently  devised  by  S.  Tashiro  in  connection  with 
his  important  researches  on  carbon  dioxide  production  in 
nerves.     By  these  methods  it  is  shown,  pointing  to  a  single 


Growth  Integration  111 

instance,  that  in  pieces  of  a  flat-worm  isolated  by  cutting 
"the  rate  of  metabolism  is  higher  in  long  anterior  pieces 
than  in  posterior  pieces  of  tlie  same  length.''  ^'^ 

Starting  from  this  low  but  seemingly  universal  level  of 
integrative  phenomena  in  tlie  individual,  Child  foniuilates 
views  of  the  nature  of  organisms  that  agree  verv  well  with 
the  organismal  standpoint  upheld  in  this  volume.  lie 
writes:  "Tlie  organic  individual  apj)ears  to  be  a  unitv  of 
some  sort.  Its  individuality  consists  primarily  in  tliis  unity, 
and  the  process  of  individuation  is  the  progress  of  integra- 
tion of  a  mere  aggregation  into  such  a  unity,  for  this  unity 
is  not  simply  the  unity  of  a  chance  aggregation,  but  one  of 
a  very  particular  kind  and  highly  constant  character  for 
each  kind  of  individual.  In  all  except  the  simplest  individ- 
uals it  determines  a  remarkable  degree  of  uniformity  and 
consistency,  both  in  the  special  relations  of  parts  and  the 
order  of  their  appearance  in  time,  and  also  in  coordination 
or  harmony  of  functional  relation  to  these  parts  after  their 
development."  ^^ 

Meristic  Gradients  and  Metabolic  Gradients  Both 
Phenomena  of  Groieth  Integration 

In  view,  then,  of  the  exceedingly  wide  ])revalence  in  living 
nature  of  axially  disposed  meristic  series  quantitivcly 
graded,  and  of  the  equally  wide  or  even  wider  })revalence  of 
axial  gradients  on  the  basis  of  metabolic  activity,  the 
gradients  of  both  sorts  arising  as-  fundamental  growtli  plie- 
nomena,  it  appears  im])ossible  to  avoid  recognizing  our  first 
category  of  integration,  namely,  groxcth  integration,  as 
a})out  the  most  simple  and  ])rimal  and  universal  of  all  these 
categories,  at  least  for  nudticellular  organisms.  It  seems 
as  though  the  other  kinds  of  differentiation  and  integration 
are  superposed,  as  one  might  express  it,  upon  this  primordial 
kind.      To  a  consideration  of  the  other,  su])erimpos('d   inte- 


112 


The  Unity  of  the  Organism 


grations  we  now  pass,  taking  them  again  in  their  seeming 
order  of  obviousness. 


REFERENCE  INDEX 


1.  Torrey   139 

Q.  Ritter    ('09)    71 

3.  Robertson  612 

4.  Moeser     373 

5.  Child   (2)    65 

6.  Child   (1)    213 


7.  Child  (1)  215 

8.  ChUd  (1)  

9.  Child  (1) 

10.  Child  (1)  

11.  Child  (2)  


66 
77 
73 

2 


Chapter  XVlll 
CHExAIICO-FUNCTIONAI.  INTEGRATION 

Functional  as  Contrasted  with  Growth  Integration 

\  SSUMING  Child's  theory  of  metabolic  gradients  to  be 
^  *"well  grounded,  we  are  furnished  thereby  with  one  im- 
portant insight  into  the  chemical  processes  involved  in  the 
unit}^  of  the  individual  organism.  But  that  process  is  con- 
cerned primarily  with  the  growth,  with  the  production  of  the 
individual.  The  question  now  is,  are  there  chemical  proc- 
esses the  object  of  which  is  to  maintain  the  functional  unity 
of  the  complete  or  nearly  complete  individual.''  Are  there 
chemical  operations  the  office  of  which  is  to  preserve  a 
proper  interrelation  among  the  parts  of  the  organism  as 
these  perform  their  special  offices.'^ 

That  such  is  to  some  extent  the  significance  of  most  if 
not  all  internal  secretions  as  usually  understood  is  indicated 
by  the  fact  that  the  functional  disturbances  attending  re- 
moval of  the  thyroids  or  other  glands  from  various  animals ; 
and  by  the  further  fact  that  where  internal  secretions  play 
a  part  in  development,  their  action  is  rather  tliat  of  stinuila- 
tor,  or  at  least  modifier,  than  of  true  producer. 

The  conception  of  internal  secretions  as  being  at  Kast  as 
much  regulators  of  physiological  function  as  of  growth  is 
illustrated  by  cases  of  hypopituitarism  of  the  post- 
adolescent  type,  like  those  described  by  Gushing,  for  ex- 
ample. In  the  series  of  cases  of  disease  due  to  "jiituitary 
deficiency"  the  first  symptoms  appeared  when  tlie  subjects 
were  from  thirty  to  forty  years  old. 

113 


114  The  Unify  of  the  Organism 

The  Conception  of  ''Internal  Secretions" 

The  nature  of  the  phenomena  now  to  be  considered,  and 
their  significance  for  our  discussion  make  it  desirable  to 
think  about  these  secretions  from  tlie  broad  standpoint  first 
stated,  according*  to  Ba3^1iss,  by  Brown-Sequard  and  d'Arson- 
val,  namely,  as  materials  produced  by  any  living  cells  or 
tissues  wliich  are  discharged  into  the  blood  or  lymph  and 
have  specific  effects  on  other  parts  or  functions  of  the  or- 
ganism. Regarded  thus  it  is  now  known  that  many  cells  of 
the  organism  produce  internal  secretions.  Although  we  are 
more  concerned  with  function  than  witli  structure  in  this 
discussion,  our  purpose  will  be  best  served  by  beginning 
with  a  mor2:)hological  classification  of  the  secretion-pro- 
ducing cells.  They  may  be  divided  into  two  categories, 
those  which  are  disposed  into  definite  organs,  or  glands,  the 
ductless  glands  of  long  standing  in  anatomy ;  and  those 
which  are  not  assembled  in  such  organs.  Knowledge  of  this 
second  class  of  cells  is  of  recent  date,  and  is  fuller  from  the 
functional  than  from  the  structural  standpoint.  The  chief 
glands,  to  which  the  name  Endocrine  has  lately  been  given 
by  Schafer,  are  now  so  well  known  as  hardly  to  need  men- 
tion. They  are  the  thyroid  apparatus,  including  the  thy- 
roids and  the  parathyroids,  the  suprarenal  body,  the  pitui- 
tary body,  and  probably  the  thymus  and  pineal  bodies.  Cells 
now  known  to  produce  internal  secretions  but  which  are  not 
arranged  in  glands  are  certain  cells  of  the  pancreas  scat- 
tered among  the  pancreatic  cells  proper ;  certain  cells  of 
the  alimentary  mucous  membrane;  the  interstitial  cells  of 
the  ovary  and  of  the  testis,  and  probably  certain  cells  of 
the  placenta,  of  the  mammary  gland,  and  of  the  uterus. 

Following  our  usual  course  of  making  the  treatment 
merely  illustrative  rather  than  aiming  at  exhaustiveness, 
our  selection  will  include  one  example  from  each  of  these 
groups.     From  the  glandular  category  we  take  the  thyroid 


Chemico-Functional  Inteyration  115 

apparatus,  and  from  tlie  non-glandular  a  jxjrtion  of  the 
alimentary  nuicous  membrane,  namely,  that  of  the  duo- 
denum. 


Effects    of    liemoving    the    Human    TliifroUl   for    Curative 

Purposes 

As  definite  knowled^-e  of  the  ^'reat  [)hyslolo^ieal  impor- 
tance of  internal  secretions  beg-ins  with  human  surgery — 
with  operations  on  the  thyroid  aj)paratus — we  may  well 
begin  our  study  here.  This  is  the  better  starting  })oint 
in  that  there  is  no  more  striking  illustration  of  how  great 
a  part  of  the  whole  organism  ma}'  be  implicated  in  the 
action  of  internal  secretions  than  is  afforded  by  the  jjrod- 
ucts  of  the  thyroids  and  parathyroids. 

The  subject  first  came  into  clear  lif^ht  in  the  early  eii^hties 
of  the  last  century  through  the  experiences  of  Swiss  sur- 
geons, Theodor  Kocher  and  J.  L.  Reverdin  esj>ecially,  who 
removed  the  thyroids  to  cure  goitre,  this  disease  being  s})e- 
cially  prevalent  in  some  parts  of  Switzerland.  The  })atients 
operated  on  were  found  to  improve  ra])idly  for  a  time  after 
the  operation,  but  later  untoward  symptoms  began  to  mani- 
fest themselves.  Because  the  variety  and  ]X'rvasiveness  of 
these  symptoms  in  a  typical  case  are  highly  instructive  for 
us  we  present  them  in  detail,  selecting  a  description  from 
Human  Physiology  by  Luciani:  "Patients  who  have  under- 
gone total  thyroidectomy  .  .  .  experience  the  initial  symp- 
toms of  glandular  deficiency  either  at  once  or  at  latest  some 
weeks  after  the  operation.  They  feel  weak,  com])laiii  of 
heaviness  of  the  limbs,  and  more  or  less  diffuse  dull  pains, 
])articularly  in  the  legs,  which  may  iH'come  acute  and  assume 
tlic  character  of  ])airis  in   the  bones. 

"Other  more  serious  symptoms  arc  gradually  associati'd 
with  the  preceding.  After  four  or  five  months  the  face 
and  the  extremities  swell  and  become  cold,   the  nmscles  are 


116  The  Unity  of  the  Organism 

torpid,  sometimes  rigid,  often  exhibiting  muscular  tremors, 
and  are  incapable  of  carrying  out  any  delicate  manual  acts 
of  precision.  At  first  the  swelling  is  variable;  it  is  more 
pronounced  in  the  morning  than  in  the  evening,  but  steadily 
increases  until  it  becomes  permanent.  It  is  not  ordinary 
oedema,  in  which  percussion  with  the  fingers  leaves  a  depres- 
sion ;  it  is  a  hard  and  elastic  swelling.  It  is  specially  local- 
ized in  the  hands,  feet  and  face,  where  it  produces  a  char- 
acteristic alteration  of  the  countenance.  The  lower  eyelids 
are  the  first  to  present  a  sacculated,  semi-transparent  swell- 
ing, which  is  hard  to  the  touch ;  then  the  infiltration  spreads 
to  the  folds  of  the  face,  which  become  smoothed  out ;  to  the 
nose,  which  gets  rounded;  to  the  lips  which  swell,  and 
bulge  outward,  saliva  dribbling  from  tliem.  The  features 
are  coarsened  and  expressionless  like  those  of  a  cretin. 

"The  mental  functions  accord  with  this  appearance,  since 
they  are  blunted,  so  that  the  patients  lose  their  memory, 
become  deaf,  taciturn,  melancholy,  self-absorbed,  and  reply 
extremely  slowly  to  questions.  They  further  complain  of 
slight  but  perpetual  headache ;  feel  an  almost  constant  sen- 
sation of  cold,  which  is  most  acute  at  tlie  extremities ;  at 
times  they  are  seized  with  vertigo,  and  may  even  lose  con- 
sciousness. 

"All  the  symptoms  become  still  further  aggravated.  The 
whole  body  may  grow  more  bulky  from  the  extension  of  the 
swelling.  The  skin  loses  its  elasticity,  can  only  be  picked 
up  in  large  folds,  and  becomes  dry  owing  to  defective 
capacity  for  sweating.  The  epidermis  desquamates  in  more 
or  less  extensive  lamellae,  particularly  on  the  hands  and  feet ; 
the  hair  turns  grey,  falls  out,  and  gets  constantly  thinner. 

"The  heart  functions  weakly,  but  with  ordinary  rhythm; 
the  pulse  is  small  and  thready.  Examination  of  the  blood 
shows  nothing  constant ;  but  there  is  often  a  more  or  less 
pronounced  and  progressive  oligocythsemia,  which  undoubt- 
edly  contributes   to   the   characteristic   pallor   of   the   skin, 


Chemico-Functional  Integration  117 

this  being  of  the  earthy,  yellow-spotted  hue  peculiar  to 
cretins. 

"The  respiratory  rhythm  is  ahnost  ahvays  normal ;  tlie 
digestive  apparatus  functions  well,  as  also  the  urinary 
system.     The  spleen  is   not  enlarged."  ^ 

This  complex  (syndrome  in  medical  terminology)  of 
manifestations    is    known    technically    as    cachexia    thyreo- 

priva. 

Later  experience  by  other  surgeons  with  the  same  opera- 
tion discovered  that  in  some  cases  the  effects  are  much  more 
acute  and  rapid,  and  may  be  replaced  by  what  has  been 
called  "tetany"  (though  having  little  in  common  with  ordi- 
nary tetanus),  ending  in  death  more  often  than  otherwise. 

Experimental  Thyroid  Excision  in  Normal  Lower  Animals 

No  sooner  had  the  far-reaching  influence  of  the  thyroid 
for  tlie  human  organism  begun  to  be  recognized  in  this  way 
than  experimentation  on  inferior  mammals  was  invoked  for 
further  light  on  the  subject.  Moritz  Schiff,  the  ground- 
breaker  in  this  field,  published  in  188-i  the  results  of  the 
removal  of  the  thyroid  from  a  large  number  of  dogs.  In 
all  cases  where  the  whole  thyroid  apparatus  was  excised 
the  dogs  soon  died  after  a  run  of  such  symptoms  as  tremor, 
spasms,  and  convulsions.  Nor  did  SchifF  rest  content  with 
merely  ascertaining  the  effects  of  removal,  complete  and 
partial,  of  the  thyroid  apparatus.  He  found  that  these 
effects  could  be  entirely  prevented  by  grafting  a  portion  of 
the  gland  under  the  skin  or  into  the  body  cavity  of  tlu' 
animal  before  the  thyroid  operation,  or  by  injecting  thyroid 
juice  into  the  blood  or  lymphatic  systems,  or  by  feeding  raw 
thyroid  to  the  dogs.  The  story  of  how  these  experiments 
led  to  the  now  widely  practiced  treatment  of  myxo^lema  with 
thyroid  or  thyroid  extract  would  be  out  of  ])lace  here, 
though  it  should  not  be  passed  wholly  unnoticed. 


118  The  Unity  of  the  Organism 

So  an  enormous  mass  of  evidence,  experimental,  surgical 
and  clinical,  is  now  in  court  demonstrating  that  for  some 
animals  at  least,  among  them  being  the  human  and  the 
canine  species,  products  of  the  thyroid  apparatus  are  in- 
dispensable to  the  normal  life,  the  symmetrical  growth  and 
balanced  physiological  activities  of  the  organism.  That 
the  apparatus  is  essential  to  tlie  "Hormonic  Equilibrium" 
of  the  organism  in  some  animals  is  beyond  question. 

While  no  pretense  can  be  made  at  an  exhaustive  exami- 
nation of  this  evidence  two  phases  of  our  discussion  make 
it  desirable  to  cary  the  examination  on  the  manifestational 
side  a  little  farther.  One  of  these  is  the  importance  of 
making  as  objective  and  emphatic  as  possible  the  extent  of 
the  manifestations  in  the  individual ;  the  other  is  the  ques- 
tion of  the  generality,  taxonomically  speaking,  of  the  thy- 
roid apparatus. 

In  the  interest  of  the  first  of  these  I  present,  verbatim,  the 
report  of  a  single  case  of  complete  thyroidectomy,  the  animal  in 
this  instance  being  a  fox.  The  individual  concerned  was  a  female 
less  than  one  year  old. 

"Oct.  28.     Glands  removed;  good  recovery. 

"Oct.   29-      Normal,  but  does  not  eat. 

"Oct.  30.  Salivation,  rapid  breathing,  strong  tremors  and  tet- 
any from  7  A.  M.  to  2  P.  M.;  quiescent  but  weak  during  the  rest 
of  the  afternoon. 

"Nov.  1.  Normal,  but  rather  weak;  eats;  no  sign  of  tremors 
or  salivation  during  the  day. 

"Nov.  2.      Restless;  slight  tremors;  dyspnoea;  does  not  eat. 

"Nov.  3.  Some  depression,  but  no  tremors  or  salivation  until 
4  P.  M. ;  does  not  eat.  At  4  P.  M.  spasms  appeared  and  con- 
tinued unabated  as  long  as  observed  (7  P.  M.). 

"Nov.  4.  Found  dead  at  4  A.  M.  Post-mortem  examination 
revealed  no  parathyroids  nor  accessory  thyroids."  ^ 

As  to  taxonomic  range  and  character  of  manifestation  of  thy- 
roid influence,  much  diversity  might  have  been  anticipated  on 
general  natural  history  grounds.  As  far  as  investigations  have 
gone  they  realize  these  anticipations.  A  summary  of  results  will 
serve  our  purpose,  and  this  is  at  hand  in  Schafer's  volume  already 


Chemico-Fujicfional  Integration  119 

cited. 

Concerning  the  effects  of  removal  of  tlie  tliyroid  appa- 
ratus he  says : 

"The  most  acute  symptoms  are  exliihited  hy  carnivora  such  as 
dogs^  cats,  foxes^  and  wolves  (Vincent),  and  tlie  young  of  lierhi- 
vora  (v.  Eiselsberg,  Sutherland  Simpson)  and  are  of  a  nervous 
nature.  .  .  .  Some  species  exliibit  no  symptoms  whatever — at 
least  when  the  operation  is  })erfornu'd  on  the  adult.  Horslcy 
states  that  this  is  the  case  with  birds  and  rabbits;  but  according 
to  Gley,  the  latter  are  affected  if  care  is  taken  to  find  and  re- 
move all  four  parathyroids,  and  Dp3'on  and  ,Jouty  obtained  tyi)ical 
tetany  in  hens  which  had  been  parathyroidectomized.  .  .  .  Ac- 
cording to  Vincent  and  Jolly  badgers  are  totally  unaffected  by 
complete  removal  of  both  thyroids  and  parathyroids."  ^ 

From  the  anatomical  characteristics  of  the  organs, 
and  from  tlie  known  effectiveness  of  minute  portions  of 
them,  such  statements  as  the  last  must  be  taken  with  re- 
serve. Although  these  results  show  by  their  diversity  that 
an  enormous  amount  of  study  remains  to  be  done  on  the 
comparative  side,  they  leave  no  question  that  the  secretion 
of  the  thyroid  af)paratus  is  important  for  the  general 
health  and  equilibrium  of  most  animals  in  which  it  occurs. 
The  measure  of  tliis  importance  in  the  eyes  of  some  authori- 
ties is  seen  in  sucli  a  statement  as,  "No  cell  anywhere  in  the 
body  can  reach  morphological  perfection  without  thyroid 
stimulus."  * 


The  Internal  Secretion  of  the  Duodenal  Mucous  Mcnihraue 

We  now  pass  to  an  examination  of  thi-  effects  of  the 
internal  secretion  of  the  duodenal  nuicons  lucnibrafu".  'I'his 
particular  secretion  is  selected  for  the  n-asons  that  it  is, 
according  to  Bayliss,  one  of  its  discoverers,  "the  most 
typical  of  all  the  chemical  messengers'';  that  it  was  om-  of 
the  first  to  be  investigated;  and  that  it  is  one  of  the  Uw 
which  have  been  isolated  as  definite  substances. 


IW  The  Unity  of  the  Organism 

The  mode  of  operation  of  this  secretion  is  tersely  stated 
by  Bayliss :  "Food  entering  the  duodenum  causes  the  pro- 
duction of  a  special  substance  which  enters  the  blood  and 
excites  the  pancreas  to  pour  into  the  duodenum  a  digestive 
juice.    ** 

That  the  presence  of  various  substances,  especially  acids, 
in  the  duodenum,  induces  a  flow  of  pancreatic  juice  was 
known  when  Bayliss  and  Starling  began  their  work  in  this 
field;  but  up  to  that  time  the  excitation  of  the  pancreas  to 
such  action  was  supposed  to  be  through  a  nerve  reflex. 
These  investigators  had  reported  in  1902  ^  that  acid  in  the 
duodenum  is  able  to  cause  the  pancreas  to  secrete  after 
nervous  communication  between  the  intestinal  wall  and  the 
pancreas  is  excluded.  They  went  further  and  obtained  an 
extract  from  the  duodenal  mucous  membrane  which,  being 
injected  into  a  vein,  induced  a  copious  flow  of  pancreatic 
juice.  The  substance,  whatever  it  is,  which  acts  thus  they 
call  secretin.  It  has  been  surmised  by  a  few  physiologists 
that  the  eff'ects  are  not  due  to  the  direct  action  of  the  secre- 
tion on  the  pancreatic  gland-cells,  but  that  the  influence 
is  exerted  through  the  vaso-dilator  mechanism.  But  this 
surmise  is  negatived  by  the  demonstration  that  the  secretin 
will  induce  the  flow  of  pancreatic  juice  while  it  does  not 
alter  the  blood  pressure.  The  case  seems,  then,  fully  estab- 
lished, and  is  so  clear-cut  and  relatively  simple  an  instance 
of  the  coordinated  functioning  of  two  wholly  distinct  parts 
of  the  body  through  chemical  means,  that  it  is  desirable 
to  get  sharply  before  us  the  known  steps  in  the  process. 
In  the  course  of  normal  digestion,  food  acidulated  in  the 
stomach  passes  into  the  duodenum.  Here,  probably  in 
virtue  of  its  acidity,  it  acts  upon  the  cells  of  the  mucous 
membrane  in  sucli  a  way  as  to  induce  them  to  produce  a 
substance  which  is  discharged,  not  into  the  intestines,  there 
to  take  its  part  in  digestion,  but  into  the  blood.  By  the 
blood   stream   the   substance   is    carried   through   its   whole 


Chemico-Functional  Integration  121 

circuit,  lience  tlirough  the  lungs  and  so  on,  around  to  the 
pancreas,  the  typical  gland-cells  of  which  it  excites  into 
activity,  so  that  the  pancreatic  juice,  an  "external"  instead 
of  an  ''internal"  secretion,  is  poured  into  the  duodenum  to 
exercise  its  digestive  office  on  the  same  food  which  started 
the  cycle  of  activities. 

It  was  with  tliis  substance  particularly  before  tlieir  minds 
Hiat  tlic  authors  adopted  the  name  Iwnnone  to  designate 
substances  which  act  tlius.  "The  group  of  substances  re- 
ferred to,"  says  Bayliss,  "which  includes  adrenaline  and  the 
various  internal  secretions,  is  characterized  by  the  prop- 
erty of  serving  as  chemical  messengers,  by  wliicli  the  activity 
of  certain  organs  is  coordinated  with  that  of  others.  They 
enable  a  chemical  correlation  of  the  functions  of  the  organ- 
ism to  be  brought  about  through  the  blood,  side  by  side 
with  that  which  is  the  function   of  the  nervous   system."  ^ 

Tliis  reference  to  the  side-by-side  activities  of  chemical 
messengers  and  nervous  system  in  integrating  the  organism 
touches  a  subject  of  the  utmost  importance.  Considera- 
tion of  it  must,  however,  be  deferred  until  we  have  looked 
a  little  more  into  the  nature  of  hormones. 

The  Nature  of  the  Actii*e  Substances  in  Internal  Secretions 

That  the  peculiar  iodine-rich  albuminous  substance  ob- 
tained from  the  thyroid  by  Baumann  in  1895  and  since 
observed  by  other  investigators,  contributes  in  some  essen- 
tial way  to  the  action  of  the  secretion  of  tliis  gland  is  the 
belief  of  apparently  a  large  majority  of  autliors  (Bayliss, 
Kppinger,  Howell,  etc.),  but  not  of  all  (Luciani).  In  view 
of  the  uncertainty  on  the  point  Schafer's  proposal  "to  ex- 
press our  ignorance  by  a  term  which  implies  no  theory" 
may  well  be  accepted,  with  the  proviso  that  the  term  pro- 
posed be  really  taken  as  evidence  that  sowefhing  though 
not  everything  is  known   about   the  substance.      A   part   of 


122  The  Unity  of  the  Organism 

the  proposal  is  worth  quoting.  "I  propose  therefore  pro- 
visionally to  apply  the  word  thyrine  to  denote  the  active 
principle,  whether  it  be  identical  with  or  contained  in  the 
iodothyrin  of  Baumann  or  not."  ^ 

Thyrine  then  becomes  the  name  of  a  substance  the  source 
and  some  of  the  activities  of  wliich  are  known,  but  whose 
main  physical  and  chemical  attributes  are  unknown. 

Concerning  the  mode  of  action  of  thyrine  there  are  sev- 
eral divergent  views,  all  based  on  some  evidence  and  so 
perhaps  not  entirely  antagonistic.  Is  the  antitoxic  theory 
of  Luciani  ^  partly  right,  right  as  regards  the  parathyroid 
secretion  (Moussu,  Vassale  and  Generali),  and  partly 
wrong,  wrong  as  to  the  secretion  of  the  thyroid  proper,  this 
being  trophic  rather  than  antitoxic?  ^^  ^lay  there  not  be 
more  in  the  enzymic  theory  suggested  some  years  ago,  than 
later  writings  have  been  inclined  to  favor  .^  Does  the  fact 
that  internal  secretions  seem  to  be  simpler  than  enzymes, 
as  indicated  by  their  greater  resistance  to  heat,  preclude  the 
possibility  that  their  normal  mode  of  action  is  of  the  enzyme 
type  after  all.?  That  is,  may  it  not  be  necessary  to  extend 
the  conception  of  enzymic  action  (which  is  surely  generic 
anyway)  to  include  the  various  sorts  of  activity  presented 
by  hormones,  understood  in  the  sense  given  it  by  its  origi- 
nators ? 

But  neither  can  the  resemblance  of  internal  secretions 
to  drugs,  so  far  as  their  action  is  concerned,  be  overlooked. 
This  has  been  dwelt  upon  by  Scliafer  ^^  and  has  important 
bearings  on  the  problems  of  the  origin  as  well  as  on  the 
chemical  nature  of  the  substances. 

Another  aspect  of  the  mode  of  action  of  internal  secre- 
tions is  that  of  whether  the  effects  are  to  stimulate  or  inhibit 
the  activity  of  the  organ  or  tissue  on  which  they  operate. 
Schafer  and  others  make  a  special  point  of  this,  directing 
the  attention  to  the  fact,  by  way  of  illustration,  that  the 
adrenaline  of  the  suprarenal  medulla  causes  contraction  of 


Chemico-Functional  Integration  123 

the  plain  muscle  of  the  blood  vessels  and  inhibition  of  that 
of  the  intestines. ^^  The  distinction  has  an  undoubted  natu- 
ral grounding',  and  so  is  in  tlio  interest  of  acenrate  doscriji- 
tion  and  clear  conce})tion. 

As  to  the  actual  chemical  composition  of  internal  secre- 
tions, knowledge  is  exceechngly  meager.  More  is  known 
about  adrenaline,  the  active  [)rinciple  of  the  su])rarenal 
gland,  than  about  that  of  the  secretion  of  any  other  gland 
or  tissue.  This  was  isolated  by  the  Japanese  chemist 
Jokichi  Takamine  in  1901,  and  has  since  been  more  fully 
examined  by  several  investigators,  notably  by  T.  B.  Aid  rich. 
It  is  described  as  a  micro-crystalline  substance  occurring  in 
at  least  five  crystal  forms.  Aldrich  assigns  to  it  the  em- 
pirical formula  CoHjoNOo,  this  structure  placing  it  not  far 
from  tyrosin  in  the  benzene  or  aromatic  series.  Of  special 
interest  is  the  astonishingly  minute  quantites  which  produce 
physiological  effects.  According  to  Aldrich  0. 000001  gram 
of  an  aqueous  solution  of  the  chloride  per  kilo  of  body 
weight  injected  into  the  blood  system  raises  the  blood  pres- 
sure 14  mm.  of  mercurv.^^ 

The  chemical  nature  of  Tethelin,  lately  isolated  from  the 
anterior  lobe  of  the  pituitary  of  the  ox,  has  been  studied 
bv  its  discoverer,  Robertson.  It  is  described  as  white  or 
pale  cream  colored,  readily  powdered,  highly  deliquescent, 
and  having  a  greasy  odor  and  slightly  acid  reaction  in 
aqueous  solutions.  It  contains  1.4  per  cent  of  phosphorus 
and  four  atoms  of  nitrogen  for  everj^  atom  of  phos]ihorus. 
The  phospliorus-nitrogen  content  of  the  substance  is  con- 
sidered by  Robertson  as  specially  significant,  since  this 
seems  to  ally  it  chemically  with  "j)hytin,"  a  substance  found 
in  the  rapidly  growing  ])arts  of  plants,  and  in  milk.  The 
natural  suggestion  is  that  the  growth-promoting  substances 
in  plants,  milk,  and  the  pituitary  secretion  are  chemically 
related. 


124  The  Unity  of  the  Organism 

The  Close  Resemblances  and  Interrelations  of  the  Different 

Internal  Secretions 

Even  the  meager,  merely  illustrative  examination  of  in- 
ternal secretions  we  have  been  able  to  make  brings  out  the 
close  resemblance  there  is  between  the  several  endocrine 
glands,  and  also  between  the  physiological  effects  of  the 
various  secretions.  These  resemblances  suggest  an  intimate 
organic  interrelationship  among  all  the  internal  secretion- 
producing  parts  of  the  body. 

All  investigators  in  this  field,  no  matter  to  how  restricted 
a  section  of  it  their  efforts  are  primarily  directed,  seem  to 
come  upon  the  interdependence  of  the  sources  and  activities 
of  hormones.  To  illustrate.  Gushing,  whose  central  interest 
has  been  the  hyphophysis,  is  led  to  conclude  that  experi- 
mentally induced  hyphophyseal  deficiency  works  histologi- 
cal changes  in  many  if  not  all  tlie  other  ductless  glands. 
It  is  not  sui^prising,  consequently  that  far-reaching  theories 
have  been  elaborated  on  the  basis  of  these  relationships. 

Bayliss  refers  with  approval  to  Elliott  for  the  conserva- 
tism with  which  he  sums  up  the  present  state  of  knowledge  on 
this  aspect  of  the  general  subject.  But  even  so,  features  are 
pointed  out  "which  suggest  a  common  bond" : 

"(1)  Carbohydrate  metabolism  is  influenced,  not  only  by 
the  pancreas,  but  also  by  the  thyroid  in  super-activity,  in 
acromegaly,  and  by  the  injection  of  adrenaline. 

"(2)  Growth  is  affected  by  the  testis  and  the  cortex  of 
the  suprarenals,  arrested  by  the  absence  of  the  thyroid. 

"(3)   Nervous  implications. 

"(4)  The  pituitary  becomes  hypertrophied  when  the  thy- 
roid is  removed.  Acromegaly  may  lead  to  enlargement  of 
the  thyroid."  ^^ 

At  the  other  extreme  of  what  may  be  regarded  as  legiti- 
mate scientific  theorizing,  we  have  the  views  of  Sajous,  who 
believes  research  will  finallj^  demonstrate  a  relationship  be- 


Chemico-Fimctional  Integration  125 

twcen  all  the  ductless  glands  the  combined  functioning  of 
which  dominates  most  of  the  activities,  normal  and  patho- 
logical, of  the  organism. 

Sajous'  elaborately  worked-out  theories  of  internal  se- 
cretions, especially  in  their  relation  to  disease  and  medical 
practice,  are  opposed  at  many  points  to  prevailing  opinion 
based  on  present  day  research.  Nevertheless  regarded  from 
the  standpoint  of  general  biology  there  would  seem  to  be 
much  merit  in  his  effort  on  the  one  hand  to  find  a  common 
ground  in  the  metabolic  processes  for  all  the  phenomena  at- 
tributed to  endocrinal  activity ;  and  on  the  other  hand  to 
find  a  more  consistent  morphological  and  physiological  basis 
of  definition  and  classification  of  internal  secretions  and  the 
structures  which  produce  them  than  has  yet  been  recognized. 
For  example,  whether  Sajous  is  right  or  not  in  contending 
that  the  pituitary  body  does  not  produce  an  internal  secre- 
tion, certain  it  is  tluit  the  non-glandular  structure  of  its 
posterior  part,  extracts  of  which  alone  have  slowing  effects 
on  the  heart,  is  strongly  suggestive  to  the  critical  naturalist 
that  the  inclusion  without  qualification  of  this  part  at  least 
of  the  organ  among  the  endocrine  glands  is  an  instance  of 
what  is  known  to  taxonomists  as  "lumping"  in  classification 
— a  kind  of  practice  that  advance  in  knowledge  always  finds 
to  be  inadequate  for  purposes  that  are  critical.  Sajous' 
late  summary  of  his  views  is  highly  suggestive  to  the  general 
biologist,  even  though  it  is  excessively  theoretical  in  some 
parts. 

As  far  as  a  much  interested  outsider  can  judge,  tlie  ]i res- 
ent state  of  understanding  of  the  relationships  among  the 
internal  secretions  is  set  forth  with  exceptional  judiciousness 
by  Waller.  "There  can  be  little  doubt,"  tlie  author  o])ens 
his  discussion  by  remarking,  "that  the  various  internal  secre- 
tions are  most  closely  correlated,  yet  perha])s  the  most  dif- 
ficult, and  also  the  most  fascinating  problem  of  present  day 
medicine,  is  to  assign  to  each  its  proper  and  riglit  share  of 


126  The  Unity  of  the  Organism 

importance."  ^^  This  statement,  coupled  with  the  fact  that 
one  of  the  main  objects  of  the  discussion  is  to  display  the 
many  contradictions  which  the  author's  large  experience  as 
a  i3ractitioner  has  found  in  the  action  of  the  internal  se- 
cretions, is  about  the  most  striking,  and  from  our  stand- 
Yfomi  most  significant  tiling  about  this  paper. 

As  one  illustration  of  the  agreement  of  action  of  the  se- 
cretions, or  at  least  of  the  influence  of  the  endocrine  glands, 
it  is  pointed  out  that  changes  in  calcium  metabolism  have 
been  obsers^ed  after  removal  of  the  thymus ;  in  disease  of  the 
pituitary  and  of  the  pineal  bodies ;  after  castration ;  after 
ovariotomy ;  after  removal  of  the  suprarenals,  and  after  re- 
moval of  the  thyroids  and  parathyroids. 

Of  the  numerous  instances  of  contradiction  which  he 
brings  out,  we  mention  only  that  concerning  tetany.  This 
he  shows  may  result  from  either  removal  of  the  thyroid  or 
from  an  overdose  of  thyroid  extract.  The  explanation  of 
the  contradictions  in  the  action  of  a  mven  secretion  favored 
by  the  author  is  that  of  the  "varying  influence  of  the  other 
internal  secretions."  But  the  descriptions  given  seem  to 
leave  no  doubt  that  diff'erence  in  type  of  individuals  also 
comes  into  the  explanation.  Thus  among  children  afflicted 
with  enlarged  tonsils  and  adenoids,  the  two  distinct  types 
dependent  upon  the  character  of  the  s^^mptoms,  is  a  case  in 
point.  One  type  is  dull  and  stupid,  stunted  in  growth,  has 
dry  coarse  skin,  and  may  display  symptoms  of  rickets.  The 
other  type  is  vivacious  pliysically  and  mentally,  given  to 
peevishness,  irritability  and  quick  fatigue,  and  always  want- 
ing a  change  of  activity.  This  type  is  over-tall  for  its  age, 
perspires  readily  and  is  fine-skinned.  Both  types  of  cases 
are  benefited  at  least  for  a  time.  Waller  says,  by  treatment 
with  thyroid  extract. 

Concerning  the  general  nature  of  the  interdependence 
among  internal  secretions,  this  author's  views  seem  to  me  so 
eminently  sound  that  I  cannot  refrain  from  quoting  them  in 


CJiemico-Functional  Integration  127 

some  fullness:  "Considerable  stress  has  been  laid  upon  the 
antagonism  of  different  internal  secretions  by  various  au- 
thors. I  believe  we  should  gain  a  truer  insight  into  their 
working  if  we  dwelt  rather  upon  their  harmony.  It  does  not 
strike  me  as  a  very  high  conception  of  the  human  organism 
that  health  should  consist  in  the  balance  of  dissentient  or 
antagonistic  forces.  It  would  seem  far  more  ideal  that  all 
the  internal  secretions  should  work  together  for  the  common 
good  of  the  organism,  and  that  wlicn  some  special  demand  is 
made  upon  a  particular  gland  the  others  will  work  in  har- 
mony with  it.  Every  gland  is  probably  necessary  for  the 
perfect  activity  of  the  rest,  and  the  harmony  between  the 
glands  is  demonstrated  by  physiological  experiments.  .  .  . 
When  it  is  found  that  the  removal  of  an  organ  constantly 
induces  either  atrophy  or  hypertrophy  of  some  other  organ, 
we  can  reasonably  deduce  that  in  the  first  case  the  organ 
removed  is  essential  to  tlie  welfare  of  the  one  that  atrophies 
in  its  absence,  and  in  the  second  case  that  the  hypertroph^'ing 
organ  is  endeavoring  to  replace  the  lost  one,  in  some  de- 
gree, and  that  therefore  the  two  organs  have  a  kindred 
function."  ^^ 

As  an  example  of  the  first  case,  the  fact  is  cited  that  the 
removal  of  the  thyroid  or  of  the  anterior  part  of  the  pitui- 
tary induces  the  atrophy  of  the  testicles  or  the  ovaries.  The 
second  case  is  illustrated  by  the  hypertrophy  of  tlie  supra- 
renal from  the  removal  of  the  thvroids,  and  also  bv  the 
hypertrophy  of  either  the  thyroid  or  tlie  pituitary  on  re- 
moval of  the  other.  "The  demonstrated  facts  of  liyper- 
trophy,"  we  read,  "clearly  point  to  an  entente  or  even  a 
triple  alliance  between  thyroid,  hy]'>ophysis  and  suprarenals. 
And  the  genital  system  is  absolutely  deix^ndent  upon  the 
integrity  of  these  three."  ^^ 

Stating  now,  in  a  single  paragra])li,  the  results  of  inves- 
tigations in  this  field,  we  have:  The  different  parts  and  ac- 
tivities of  the  organism  are  maintained  in  their  normal  state, 


128  The  Unity  of  the  Organism 

both  as  to  the  essential  nature  of  each,  considered  individ- 
ually, and  as  to  their  relation  with  one  another,  by  a  number 
of  exceedingly  powerful  and  subtle  chemical  substances  (in- 
ternal secretions)  which  are  produced  by  certain  parts,  are 
passed  into  the  blood,  and  by  it  are  carried  about  over  the 
whole  organism  to  exert  their  appropriate  influences  on 
other  parts  and  functions.  Because  of  the  peculiar  way  these 
substances  do  their  work,  they  have  been  called  chemical 
messengers,  or  to  have  a  distinctive  name,  hormones. 

Relation  Between  the  Internal  Secretory  and  Nervous 

Systems 

But  no  physiological  truth  is  better  known  than  that  one 
of  the  main  offices  of  the  nervous  system  is  to  correlate  the 
organs  and  parts  of  the  body  with  one  another.  It  is  but 
natural  to  suppose,  therefore,  that  if  there  is  a  chemical 
scheme  for  accomplishing  the  same  end,  the  two  are  in  some 
way  closely  related. 

That  the  relations  which  exist  between  the  cerebro-spinal 
nervous  system,  the  autonomic  nervous  system  (including  the 
sympathetic),  and  the  internal  secretions,  constitute  one  of 
the  most  important  subjects  in  the  whole  physiological  do- 
main, at  the  same  time  that  it  is  one  of  the  most  recondite 
and  difficult  to  investigate,  has  come  gradually  to  view 
through  the  work  of  the  last  few  decades.  We  will  try  to 
extract  enough  from  the  mass  that  has  been  written  on  the 
subject,  to  illustrate  the  principles  involved.  The  modern 
period  of  knowledge  of  what  was  formerly  but  rather  in- 
definitely included  under  the  term  sympathetic  nervous  sys- 
tem, has  revealed  that  we  have  to  do  with  a  portion  of  the 
general  nervous  mechanism  which  in  reality  is  a  subdivision 
of  a  larger  category. 


Chemico-Functioiial  Integration  129 

Composition  and  Nature  of  the  Autonomic  System 

The  name  autonomic  was  given  to  this  category  by  Lang- 
ley.  "The  autonomic  nervous  system,"  he  says,  "means  the 
nervous  system  of  the  glands  and  of  the  invohuitary  muscles ; 
it  governs  the  'organic'  functions  of  the  body";  and  further: 
"The  word  implies  a  certain  degree  of  independent  action, 
but  exercised  under  the  control  of  a  higher  power."  ^**  l^ay- 
liss  adds:  "It  is  necessar}^  to  be  quite  clear  that  the  au- 
tonomic system  includes  the  sym]:)athetic,  since  some  writers 
abroad  use  the  name  as  applying  to  all  the  visceral  nervous 
system  other  than  the  sympathetic,  speaking  of  sympathetic 
and  autonomic."  ^^ 

Perhaps  the  most  important  single  fact  which  differen- 
tiates the  autonomic  from  the  cerebro-spinal  system  is  the 
intercalation,  everywhere  in  the  autonomic  system,  of  an 
extra  neurone  between  the  cerebro-spinal  ners'e  and  the  part 
innervated.  Cannon  states  this  distinction  very  clearly : 
"The  skeletal  muscles  receive  their  nen-e  supply  direct  from 
the  central  nervous  system,  i.  e.,  the  nerve  fibers  distributed 
to  these  muscles  are  parts  of  the  neurones  whose  cell-bodies 
lie  within  the  brain  or  spinal  cord.  The  glands  and  smootli 
muscles  of  the  viscera,  on  the  contrary,  are,  so  far  as  is  now 
known,  never  innervated  directly  from  the  central  nervous 
s^'stem.  The  neurones  reaching  out  from  the  brain  or  spinal 
cord  never  come  into  immediate  relation  with  the  gland  or 
smooth  muscle  cells  ;  there  are  always  interposed  between  the 
cerebrospinal  neurones  and  the  visceral  extra  neurones 
whose  bodies  lie  wholly  outside  the  central  nervous 
system."  ^^ 

Cannon's  suggestion  that  these  interposed  neurones  may 
function  as  transformers  for  impulses  received  from  the  cere- 
brospinal system  should  be  noted  here. 

Three  sharp  subdivisions  of  the  autonomic  nervous  sys- 
tem are  recognizable.     One  is  known  as  the  vagal  or  cranial 


130  The  Unity  of  the  Organism 

autonomic,  because  it  is  largely  made  up  of  fibers  from  the 
vagus,  or  tenth  pair  of  cranial  nerv^es.  Another  is  the  sym- 
pathetic, or  better,  the  thoracico-lumbar  autonomic,  be- 
cause its  fibers  originate  from  the  great  visceral  sympathetic 
ganglia.  This  is  by  far  the  most  extensive  of  the  three  sub- 
divisions, and  is  the  only  one  that  is  distributed  to  all  parts 
of  the  body.  The  third  is  the  sacral  autonomic.  As  its  name 
implies,  it  is  quite  restricted  in  extent,  its  fibers  being  dis- 
tributed to  the  extreme  distal  end  of  the  intestine,  the  urin- 
ary bladder,  and  some  of  the  external  genital  organs.  But 
the  differences  between  the  three  which  are  most  important 
for  us  are  physiological,  a  particularly  important  difference 
being  that  the  thoracico-lumbar  di\'ision  acts  antagonistic- 
ally to  both  the  end  divisions.  Stimulation  of  the  fibers  of 
the  sympathetic  has  just  the  opposite  effect  to  the  same 
stimulus  applied  to  the  fibers  of  the  others.  "The  sympa- 
thetic fibers  check,  the  vagal  autonomic  fibers  excite,  the 
movements  of  the  intestines ;  the  sympathetic  dilates,  the 
vagal  autonomic  contracts,  the  pupil ;  the  sympathetic 
hastens,  the  vagal  autonomic  slows,  the  heart."  ^^  The 
sacral  contracts  the  lower  part  of  the  large  intestine  and  re- 
laxes the  outlet  of  the  bladder,  while  the  sympathetic  relaxes 
the  same  part  of  the  intestine  and  contracts  the  same  part 
of  the  bladder.  Cannon  states  the  general  principle  thus : 
''When  the  mid-part  meets  either  end  part  in  any  viscus 
their  effects  are  antagonistic.'*  ^^ 

While  the  incompleteness  of  knowledge  in  this  field  needs 
emphasizing,  yet  that  knowledge  is  sufficient  to  put  some  of 
the  main  features  beyond  question,  and  to  make  clear  the 
great  importance  of  the  subject  and  of  fuller  knowledge  on 
it.  Touching  these  general  aspects  Professor  L.  F.  Barker 
writes :  "While  we  do  not  yet  understand  the  exact  mechan- 
isms of  association  among  the  activities  of  the  cerebrum, 
the  endocrine  glands,  and  the  reciprocally  antagonistic  au- 
tonomic domains  and  their  end-organs,  we  can  begin  to  see 


Chemico-Functional  Integration  131 

the  paths  which  must  be  followed  in  order  tliat  more  exact 
knowledge  may  be  gained."  -'"^ 

Experimental  Evidence  of  Connection  Between  the  Adrenal 
Glands  and  the  Nervous  System 

Some  of  the  most  important  information  we  have  in  this 
field  is  furnished  by  Cannon  and  his  collaborators  concern- 
ing  the  secretion  of  the  adrenals  and  its  relation  to  the 
autonomic  and  central  nervous  systems.  It  had  been  proved 
before  Cannon  began  his  investigations  that  adrenin  injected 
into  the  blood  has  exactly  the  same  effect  on  certain  parts 
of  the  organism  as  does  the  sympathetic  autonomic  nerves 
with  which  the  same  parts  are  supplied,  and  that  the  effect 
of  the  secretion  is  direct  and  not  through  the  nel'^'es.  In 
other  words,  it  had  been  proved  that  the  organism  has  two 
methods  by  which  the  same  activity  of  certain  of  its  parts 
can  be  induced,  one  nervous,  the  other  chemical.  Thus  the 
dilation  of  the  pupils,  the  erection  of  hairs,  the  inhibition 
of  activities  of  the  alimentarv^  canal,  and  the  liberation  of 
sugar  from  the  liver  can  be  induced  either  through  sym- 
pathetic autonomic  centers  or  by  the  secretion  of  the  adrenal 
bodies.  This  in  itself  was  important  evidence  of  interrela- 
tion between  the  nervous  system  and  internal  secretory  sys- 
tem. But  the  experimental  researches  prosecuted  in  Can- 
non's laboratory  have  proved  that  a  connection  exists  be- 
tween the  autonomic-adrenal  phenomena  and  the  cerebro- 
spinal system  through  the  sensory  nen-es,  and  with  the 
psychic  life  of  the  animal;  and  have  shown  the  jirobable  sig- 
nificance of  the  entire  scheme  for  the  life  of  tlic  organism  as 
a  whole. 

To  be  a  little  more  specific,  they  have  proved: 
(1)   That  strong  excitation  of  sensory  nerves  stimulates 
reflexly   the   adrenal   glands    and   causes   them    to   pour   an 
increased  amount  of  adrenin  into  the  blood. 


13^  The  Unity  of  the  Organism 

(S)  That  emotional  excitement  (as  the  fright  of  a  cat  by 
a  dog)  similarly  increases  the  flow  of  adrenin. 

(8)  That  this  increase  of  adrenin  in  the  blood  may  in- 
crease the  liberation  of  sugar  from  the  liver  into  the  blood 
to  such  an  extent  as  to  make  sugar  appear  in  the  urine,  thus 
demonstrating  a  true  "emotional  gl^^cosuria." 

(4)  That  the  increased  adrenin  of  the  blood  thus  pro- 
duced is  probably  advantageous  to  the  organism  in  that  it 
enhances  its  ability  to  meet  special  stresses  that  naturally 
accompany  special  excitement,  as  of  fear,  anger,  or  pain, 
this  advantage  consisting  partly  in  augmentation  of  the 
working  energy  of  the  muscles,  probably  through  the  sugar 
delivered  to  them,  and  in  increasing  the  coagulability  of  the 
blood,  thereby  reducing  the  danger  from  bleeding  wounds. 

Summing  up  his  conclusions  as  to  utility.  Cannon  writes : 
"These  changes  in  the  body  are,  each  one  of  them,  directly 
serviceable  in  making  the  organism  more  efficient  in  the 
struggle  which  fear  or  rage  or  pain  may  involve;  for  fear 
and  rage  are  organic  preparations  for  action,  and  pain  is 
the  most  powerful  known  stimulus  to  supreme  exertion.  The 
organism  which  with  the  aid  of  increased  adrenal  secretion 
can  best  muster  its  energies,  can  best  call  forth  sugar  to 
supply  the  laboring  muscles,  can  best  lessen  fatigue,  and  can 
best  send  blood  to  the  parts  essential  in  the  run  or  the  fight 
for  life,  is  most  likely  to  sursdve."  ^^ 

But  fear  and  rage  are,  one  hardly  need  be  reminded,  in 
part  psychic  phenomena,  and  hence  inseparably  connected 
with  the  higher  centers  of  the  cerebrospinal  nervous  system. 
Though  in  the  main  reflex  and  automatic,  they  are  neverthe- 
less to  some  extent  subject  in  man  to  intelligent  control. 
Thus  the  way  is  open  for  a  measure  of  rational  understand- 
ing of  the  structural-functional  means  by  which  human  be- 
ings "tap,"  as  William  James  would  say,  and  bring  under 
direction  those  remarkable  "reservoirs"  of  ordinarily  unused 
energy   about  which  everybody  knows   something   from   his 


Chemico-Functional  Integration  133 

own  experience,  and  upon   wliieli   nobody   lias  written   more 
intelligently  than  James. 

An  excellent  beginning  has  been  made,  then,  in  the  ex- 
perimental demonstration  of  the  integration  of  the  endo- 
crinal  and  common  glandular  systems,  the  blood  circulatory 
system,  the  autonomic  and  cerebrospinal  nervous  systems, 
and  the  emotional-psj^chic  life  of  animals. 

Clinical  Evidence  of  Adrenal-Nervous  Connection 

But  important  knowledge  and  general  views  in  this  field 
are  also  coming  from  clinical  medicine,  and  pharmacology. 
A  general  presentation  of  the  results  reached  down  to 
1913  is  contained  in  Innere  Sekretion  und  Nervensystem^ 
by  H.  Eppinger  and  others.  A  particularly  significant  body 
of  evidence  coming  from  this  source  concerns  the  relation 
between  the  sympathetic  or  middle  autonomic  nervous  ap- 
paratus and  the  two  end  autonomic  systems,  the  cranial  and 
sacral.  These  two  groups  act,  it  will  be  recalled,  antagon- 
istically to  each  other.  Eppinger  and  others  have  shown 
that  the  thoracico-lumbar,  or  sympathetic,  and  cranial,  or 
vagal  systems  differ  in  susceptibility  to  stimuli  In  different 
individuals,  and  perhaps  in  the  same  individual  at  different 
times,  thus  making  the  two  groups  what  Is  called  sympa- 
theticotonic  and  vagotonic  with  reference  to  each  other,  de- 
pending on  whether  the  sympathetic  or  the  vagal  is  the  more 
susceptible  to  stimuli.  This  difference  can  be  demonstrated 
by  the  administration  of  various  drugs,  as  adrenin  and  pilo- 
carpin.  But  it  is  known,  according  to  E})pinger,  that  the 
thyroid  toxin  stimulates  both  the  sympathetic  and  the  vagal. 
From  this  it  results  that  over-stimulus  of  either  may  occur 
through  this  source,  and  go  to  the  extent  of  ]M'oducIng  the 
characteristic  symptoms  of  Basedow's  or  Graves^s  disease 
(rapid  heart  beat,  exophthalmia,  diarrhea,  etc.).  Tiiese 
symptoms    may    occur    in    varying    degree,    dej)ending    on 


134  The  Unity  of  the  Organism 

whether  the  patient  is  sympathetico-  or  vagotonic,  the  sym- 
patheticotonic  type  of  the  disease  being  characterized  by 
marked  protrusion  of  the  eyeballs,  especially  rapid  heart 
beat,  absence  of  sweats,  diarrhea,  and  disturbance  of  the 
respiration ;  while  the  vagotonic  type  is  characterized  by 
slight  protrusion  of  the  eyes  and  increase  of  heart  action, 
by  outbreaks  of  sweat,  diarrhea,  and  by  faultiness  in  the 
respiratory  rhythm. 

While  some  observers,  like  Falta,  do  not  believe  the  facts 
now  known  can  be  definitely  classed  in  this  manner,  the  ef- 
fort, justified  by  some  positive  knowledge,  has  at  least  the 
merit  of  specifying  to  some  extent  the  intricate  reciprocal 
action  between  the  thyroid  apparatus  and  the  nervous  sys- 
tem, and  also  between  the  different  portions  of  the  auto- 
nomic system;  and  to  this  extent  all  students  of  the  subject 
bear  witness.  Thus  Falta :  "In  my  opinion  everything  speaks 
for  the  fact  that  in  Basedow's  disease  the  entire  nervous 
system  is  in  a  condition  of  over-excitement  and  that  the 
pictures  presented  by  the  vegetative  nervous  system  are  as 
uncommonly  manifold  and  always  changing."  "^ 

The  indication  of  prime  importance  in  this  is  that  in  these 
antagonistic  divisions  of  the  autonomic  nervous  and  endo- 
crinal  glandular  systems,  operating  together  with  the  other 
portions  of  the  organism,  there  is  a  balancing-ofF  or  equil- 
ibrating apparatus  through  which  the  whole  complex  of  vege- 
tative functions  is  carried  on,  all  of  which  in  turn  are  con- 
nected with  the  ps3^chic  functions.  Probably-  no  better  il- 
lustration can  be  found  of  the  conception  of  the  organism  as 
fundamentally  dynamic.  According  to  this  conception  nor- 
mality, both  in  function  and  in  structure,  consists  not  in 
rigid,  invariable  activities  and  organs,  but  in  a  ceaseless  play 
of  constitutively  antagonistic  forces  and  structures.  By  this 
conception  the  whole  life  of  the  organism,  physical  and  psy- 
chical, may  be  crudely  likened  to  the  performance  of  the 
tight-rope  walker,  which  depends  on  numberless   balancing 


Cliemico-Functional  Integration  1.'3.") 

activities.  Let  the  performer  be  reallv  motionles.s  in  every 
part  for  one  instant,  and  lie  falls. 

The  treatment  of  tetany  and  its  relation  to  internal  se- 
cretions, especially  to  that  of  the  thyroid,  by  Eduard  Phleps 
in  the  work  now  under  consideration,  is  another  excellent  il- 
lustration of  how  interpretation  may  run  In  accordance  witli 
this  conce])tion  of  the  animal  organism.  The  essence  of  the 
section,  as  touching  this  question,  is  contained  in  tlie  fol- 
lowing : 

"On  the  ground  of  clinical  symptoms  authoritative  clin- 
icians like  Eulenburg,  Kahler  and  Nothnagel  explained  tet- 
anus as  a  disease  of  the  entire  nervous  system.  Later  it  was 
proved  that  this  disease  was  not  due  to  primary  organic 
changes  of  the  nervous  system,  but  to  secondary  functional 
disorders.  .  .  .  The  view  of  those  authors,  who  refer  the 
disease  to  the  sim])le  effect  of  a  substance  of  the  epithelial 
granules  acting  normally  and  continuously  on  the  whole 
nervous  system,  finds  here  a  further  development  of  that  old 
theory  because  for  them  the  clinical  picture  of  a  regular 
grouping  of  nervous  stimulus-  and  response-phenomena 
arises  from  an  impairment  of  the  close  functional  relation 
between  the  nervous  system  and  the  epithelial  granules  (Mac- 
Callum,  Chvostek  jun.,  Biedl,  Eppinger,  Falta,  Rudinger, 
Jonas,  et  al.).  In  agreement  with  these  authors  we  con- 
ceive the  action  of  the  epithelio-secretive  substance  as  that  of 
a  hormone  in  the  sense  of  Starling  and  Bayliss,  which  must 
have  its  essential  point  of  attack  on  certain  reflex  stations 
of  the  central  nervous  system."  ^^ 

Taking  cognizance,  now,  of  the  fact  that  most  if  not  all 
the  cells  known  to  produce  internal  secretions  arise  embry- 
onically  from  epithelium,  as  does  also  almost  all  nervous 
tissue,  we  have  the  suggestion  of  a  deep-seated  combination 
scheme,  chemical-and-nervous,  for  integrating  the  organism. 
The  best  investigated  example  of  what  is  here  referred  to  is 
the  suprarenal  bodies.     It  is  now  fully  established  that  the 


136  The  Unity  of  the  Organism 

inner  or  medullary  portion  of  the  organ,  the  part  which  pro- 
duces the  adrenin,  is  developed  from  the  same  neuroblastic 
mass  out  of  which  the  sympathetic  autonomic  ganglia  arise. 
Furthermore,  it  seems  bej^ond  question  that  the  so-called 
chromophil,  or  sometimes  the  adrenin  granules  in  the  chief 
cells  of  the  functioning  gland  play  a  fundamental  role.  Com- 
bining these  facts  with  the  equally  well  ascertained  facts 
that  the  cortex  of  the  gland  is  derived  from  the  same  em- 
bryonic mesoblastic  mass  which  gives  rise  to  the  genital 
glands ;  and  that  in  adults  cortical  changes  in  the  supra- 
renals  are  intimately  correlated  with  reproductive  changes, 
and  a  general  view  of  the  factual  basis  on  which  the  sugges- 
tion rests  is  before  us.  We  may,  I  think,  regard  the  sugges- 
tion not  only  as  justified  but  as  revolutionary  in  comparison 
with  any  theory  that  was  scientifically  justifiable  until  re- 
cently. The  following  further  quotation  from  Phleps  brings 
the  idea  into  still  clearer  view : 

"The  unqualified  dependence  of  the  nervous  system  on  the 
epithelial  bodies  (this  last  used  in  the  sense  of  general  physi- 
ological considerations,  and  only  by  way  of  illustration), 
and  the  absolutely  vital  significance  of  this  enables  us  to  see 
many  things  in  a  new  light.  We  learn  that  many  motor, 
sensory  and  vasomotor-trophic  functions  of  the  central  nerv- 
ous system  even  up  to  the  highest  reflex  stations  having  the 
most  complicated  cortical  functions,  are  in  constant  func- 
tional cooperation  with  organs  which  heretofore  have  not  re- 
ceived sufficient  consideration  from  this  standpoint.  The 
results  compel  a  change  of  our  views  concerning  the  con- 
stantly dominating  position  of  the  nervous  system.  We  may 
see  in  many  of  its  activities  only  the  most  manifold  inter- 
mediary roUs  between  glandidar  functions  which  are  in  the 
closest  relation  to  metabolism,  and  the  sum  total  of  all  re- 
actions which  follow  external  stimuli.^'  ^^ 


Chemico-Functional  Integration  137 

Summary  of  Present  State  of  KnowJedye  In  This  Field 

Speaking  generally,  we  may  say  that  the  trend  of  all  re- 
sults, experimental  and  clinical,  is  unquestionably  toward  a 
demonstration  of  the  closest  interaction  l)etween  the  entire 
internal  secretory  system  and  both  the  autonomic  and  cere- 
brospinal nervous  systems,  this  interaction  affVcting  the 
whole  of  both  the  growth  and  the  functioning  of  the  animal 
organism. 

But  we  must  remind  ourselves  again  how  fragmentary 
knowledge  is  in  this  great  realm.  Unanswered  questions  meet 
one  on  the  ver}'^  threshold  of  any  portion  he  enters,  \^y 
way  of  illustration,  take  the  phenomenon  of  abnormal  growth 
known  as  acromegaly.  This  malady  is  characterized,  as  tlie 
name  indicates,  by  a  "peculiar  non-congenital  hypertrophy 
of  the  upper  and  lower  extremities  and  of  the  head."  ~^  Sucli, 
according  to  Schafer,  is  the  definition  given  by  Pierre  Marie, 
who  first  fully  described  the  disease.  The  main  visible  symp- 
toms consist  in  the  enlargement  of  the  bones  of  the  head, 
hands,  feet,  chest,  etc.,  especially  in  their  terminal  portions. 
Through  such  growth  the  nose  and  lower  jaw,  especially  the 
chin,  become  protrudent.  But  the  whole  skeleton  is  more  or 
less  affected,  and  there  is  a  corresponding  over-development 
of  tlie  muscles,  the  affected  person  becoming  abnormally 
strong.  That  acromegaly  is  constantly  associated  witli  an 
abnormal  condition  of  the  hypophysis  is  recognized  by  ap- 
parently all  authorities.  Whether  the  abnormality  of  the 
gland  is  a  cause  or  only  an  accom]^animent  of  the  disease  is 
an  open  question  in  the  minds  of  some.  However,  the  view 
of  a  large  majority  is  that  such  a  causal  relation  does  exist. 
"That  the  acromegalic  skeletal  growth,"  says  Schiifir,  '*is 
produced  by  hypertrophy  and  oversecretion  (or  ])erverted 
secretion)  of  the  anterior  lobe  is  highly  probable,  both  as  the 
result  of  partial  extirpation  in  animals  and  from  the  effect  of 
operative  removal  of  the  pituitary  tumours  in  man."  ^®   The 


138  The  Unity  of  the  Organism 

particular  issue  here,  it  will  be  noticed,  is  the  vitally  im- 
portant one  of  what  might  be  called  the  functional  as 
distinguished  from  the  hereditary  cause  or  at  least  incitement 
of  growth. 

In  illustration  of  the  importance  of  understanding  the 
unification  among  these  complex  systems,  the  manifestation 
of  which  is  in  turn  dependent  upon  the  organism  as  a  whole, 
the  following  from  the  address  by  L.  F.  Barker  already  re- 
ferred to,  is  impressive. 

"In  how  far  these  sudden  and  violent  excitations  of  the 
autonomic  nervous  system  which  accompany  strong  emo- 
tions are  due  to  the  intervention  of  the  glands  of  internal 
secretion,  and  in  how  far  they  depend  upon  direct  neural 
conduction  from  the  brain,  we  are  as  yet  but  ill-informed.  I 
need  only  remind  you  of  the  vasodilation  of  the  face  in  the 
blush  of  shame,  of  the  palpitation  of  the  heart  in  joy,  of  the 
stimulation  of  the  sudoriparous  glands  which  precedes  the 
sweat  of  anxiety,  of  the  stimulation  of  the  vasoconstrictors, 
the  pupil  dilators  and  the  pilomotors  in  the  pallor,  mydriasis 
and  goose-skin  of  fright,  to  illustrate  some  of  these  violent 
autonomic  excitations."  ^^ 

The  references  here,  it  will  be  noted,  are  primarily  mani- 
festations pertaining  to  the  surface,  the  integumentary 
parts  of  the  body,  and  their  scope  is  what  especially  inter- 
ests us.  Very  nearly  the  whole  list  of  these  parts  is  in- 
volved, and  probably  a  complete  inventory  would  be  still 
more  inclusive. 

Now  notice  the  range  of  manifestations  at  a  deeper  level 
that  are  involved.  "The  balance  maintained  normally  be- 
tween the  two  antagonistic  systems  the  vagal  and  sympa- 
thetic autonomic  is  one  of  the  most  interesting  of  physio- 
logical phenomena.  Think,  for  example,  of  the  rate  of  the 
heart-beat — how  constantly  it  is  maintained  at  a  given  level 
in  each  individual  when  the  body  is  at  rest ;  the  impulses  ar- 
riving through  the  vagal  system  just  balance  those  arriving 


Chemico-Functional  Integration  139 

through  the  sympathetic  system,  so  as  to  maintain  a  rate  of 
approximately  seventy-two  beats  per  minute.  And  a  similar 
balance  is  maintained  in  other  autonomic  domains  (e.g.,  the 
pupils,  bronchial  musculature,  gastric  glands,  gastro-intes- 
tinal  muscle,  sweat  glands,  bladder  muscles,  etc.)."'^'^ 

And  Barker  tlien  calls  attention  to  the  extent  to  which  the 
normal  processes  of  the  body  depend  upon  temporary  upsets 
of  these  equilibria,  examples  of  which  are  watering  of  tlie 
mouth  at  the  smell  or  sight  of  food  which  appeals  to  the 
appetite  through  these  senses,  the  flow  of  gastric  and  pan- 
creatic juices  at  the  proper  time,  through  indirect  stimula- 
tion; the  sudden  relaxation  of  the  sphincter  and  contraction 
of  the  detrusor  of  the  bladder  in  micturition ;  the  violent 
contractions  of  all  the  muscles  concerned  in  parturition  in 
the  female,  and  so  on. 

We  may  summarize  the  results  of  this  chapter  thus : 

(1)  The  researches  of  recent  years  on  the  internal  secre- 
tory sj^stem  and  its  connection  with  the  great  subdivisions 
of  the  nervous  system,  and  with  the  blood,  muscular,  and 
reproductive  systems,  have  laid  a  solid  foundation  for  an  un- 
derstanding* of  the  chemico-functional  basis  of  tlie  animal 
organism's  unity. 

(2)  The  emotional  phase  of  the  psychic  life  of  the  ani- 
mal is  proved  to  be  in  direct  organic  connection  with  tlu.^ 
basis. 

From  these  results  there  naturally  springs  the  important 
question:  What  relation  has  human  consciousness  to  this 
same  basis .?  An  attempt  to  answer  this  question  will  be  an 
unavoidable  part  of  our  treatment  later  of  the  ])sychic  in- 
tegration of  the  organism. 

REFERENCE  INDEX 

1.  Luciani   II,  13  5.  Bayliss     707 

2.  Carlson  and  Woelfel 49  6.  Bayliss  and  Starling 325 

3.  Schiifer    (1914)     18  7.  Bayliss     706 

4.  Hertoghe    194  8.  Schafer    28 


140 


The  Unity  of  the  Organism 


9.  Luciani   II,  ^ 

10.  Luciani    II,  35 

11.  Schafer    9 

12.  Schafer    11 

13.  Aldrich    457 

14.  Bayliss     721 

15.  Waller    277 

16.  Waller    280 

17.  Waller    281 

18.  Langley     240 

19.  Bayliss     484 


20.  Cannon,  W.  B.('16)    22 

21.  Garrison    144 

Q'2.  Cannon,  W.  B.   ('16)    ...       34 

23.  Garrison    146 

24.  Cannon,  W.  B.   ('13)    ...     372 
95.  Falta 59 

26.  Eppinger,  et  al 214 

27.  Eppinger,  et  al 217 

28.  Schafer    106 

29.  Schafer     69 

30.  Garrison    146 


Chapter  AV.Y 

THE    ORGAMSMAL    SIGMFICANCi:    OF     IIIK 
INTERNAL   SECRETORY    SYSIKM 

TT  remains  now  to  consider  the  real  purpose  for  wliicli 
-■-  the  presentation  of  facts  and  views  on  internal  secre- 
tions has  been  made,  namely  that  of  showing  critically  the 
significance  of  this  secretory  system  for  the  organismal 
conception. 

General  Iiiahility  of  Element  alls  m  to  Interpret  the 

Phenomena 

There  is  perhaps  no  better  way  of  approaching  this  part 
of  our  task  than  by  noticing  the  inability  of  elementalistic 
biology  to  deal  in  a  really  intelligent  and  consistent  manner 
with  the  phenomena  in  this  field.  The  breakdown  of  bio-ele- 
mentalism  when  confronted  with  the  phenomena  of  "chemical 
messengers"  nowhere  finds  more  cogent  illustration  than  in 
the  effort  to  identify  internal  secretions  with  the  organ- 
forming  substances  hypothesized  by  Sachs  and  others. 

Although  in  what  follows  the  exposure  of  inconsistency 
and  fallacy  will  haA^c  to  be  drastic  and  may  seem  personal, 
the  truth  is  it  is  wholly  impersonal  in  spirit  and  is  directed 
at  a  system  of  bad  reasoning  born  of  what  might  he  called 
a  juvenile  metaph3^sics  of  the  living  world. 

The  objective  achievements  of  Jacques  liOcl)  and  others 
of  the  school  he  represents,  in  experimental  biology,  ment 
the  admiration  of  all  lovers  of  obser^'ational  truth.  One 
may   be,    too,   more    tolerant    of   their    faults    as    reasoners 

141 


142  The  Unity  of  the  Organism 

than  he  could  be  but  for  his  recognition  of  their  high  service 
against  all  the  traditional  forms  of  supernaturalism.  The 
real  case  against  the  school  is,  as  I  see  it,  two-fold.  First,  in 
their  zeal  to  substitute  naturalism  for  supernaturalism  they 
fail  to  notice  that  supernaturalism  is  in  its  very  essence 
finalistic,  and  they  are  led  to  imagine  they  have  attained, 
or  can  attain,  natural  explanations  that  fully  supplant  the 
old  supernatural  explanations.  This  results  in  the  con- 
version of  their  supposed  naturalism  into  something  which 
is  essentially  another  kind  of  supernaturalism.  The  second 
part  of  the  case  against  the  school  is  its  abuse  of  the  most 
common  principles  of  the  knowledge-getting  processes  in  ob- 
jective biology.  For  the  general  good  of  the  biological 
sciences  the  urgent  need  of  reformation  touching  both  as- 
pects of  the  case  has  led  me  to  examine  the  particular  in- 
stance of  Loeb's  treatment  of  internal  secretions  at  greater 
length  than  would  otherwise  be  justifiable. 

Critique  of  the  View  That  Internal  Secretions  are  ^^Formative 

Stuffs" 

Although  Loeb  is  the  only  author,  so  far  as  I  know,  who 
has  expressly  contended  for  the  identity  of  internal  secre- 
tions with  Sachs'  organ-forming  substances,  the  assumption 
is  so  accordant  with  the  spirit  of  elementalism,  and  Loeb  is 
so  typical  and  eminent  a  protagonist  of  this  philosophy, 
that  his  proposal  will  probably  find  many  adherents.  It  is 
consequently  desirable  to  see  what  there  is  in  the  effort  to 
bring  hormones  into  such  a  historical  setting. 

The  statement  of  Loeb's  Aaews  is  contained  in  "The  Or- 
ganism as  a  Whole  from  a  Ph3^sico-Chemical  Viewpoint," 
1916.  Referring  to  his  espousal  twenty-five  years  ago  of 
Sachs'  hypothesis  to  explain  heteromorphosis,  he  writes :  "At 
that  time  the  idea  of  the  existence  of  such  organ-forming 
substances  was  received  with  some  scepticism,  but  since  then 


Significance  of   the  Internal  Secret  or  ij  System     143 

so  many  proofs  for  their  existence  have  been  obtained  that 
the  idea  is  no  longer  questioned.  Such  substances  are  known 
now  under  tlie  name  of  'internal  secretions'  or  'hormones'; 
their  connection  with  the  theory  of  Sachs  was  forgotten  witli 
the  introduction  of  the  new  nomenclature."  ^ 

A  case  to  which  Loeb  makes  special  reference  as  proof 
that  internal  secretions  are  the  same  as  Sachs'  organ-form- 
ing substances  is  that  of  the  effect  of  thyroid  on  the  meta- 
moi-phosis  of  the  tadpoles  of  frogs  and  toads,  demonstrated 
by  Gudernatsch.  The  author's  mode  of  using  this  case  in 
illustration  of  his  contention  is  highly  instructive.  He  re- 
fers to  the  legless  condition  of  the  tadpoles  and  calls  atten- 
tion to  so  much  of  Gudernatsch's  results  as  pertain  to  these 
members.  Gudernatsch  found,  Loeb  points  out,  that  where- 
as in  the  usual  course  of  things  the  tadpoles  live  from  four 
months  to  a  year  before  the  legs  grow  out,  by  feeding  them 
on  thyroid  gland  these  members  can  be  induced  to  appear 
at  any  time.  "We  must,  therefore,  draw  the  conclusion," 
Loeb  says,  "that  the  normal  outgrowth  of  legs  in  a  tadpole 
is  due  to  the  presence  in  the  body  of  substances  similar  to 
the  thyroid  in  their  action  (it  may  possibly  be  thyroid  sub- 
stance) which  are  either  formed  in  the  body  or  taken  up  in 
the  food."  "  When  the  case  is  presented  in  this  way  and 
nothing  more  said  about  it,  it  certainly  looks  considerably 
as  if  the  thyroid  substance  or  something  like  it  is  leg-form- 
ing substance ;  and  such  an  interpretation  would  be  enticing 
were  it  really  true,  as  Loeb  says,  that  no  other  substance 
seems  to  have  sucli  an  effect. 

When,  however,  one  consults  the  account  given  by  Gudir- 
natsch  himself  as  to  wliat  his  experiments  were  and  what 
they  established,  the  whole  matter  stands  in  a  quite  different 
light.  First  of  all,  the  fact  that  in  tliese  experiments  mam- 
malian thyroid  was  largely  used  as  food  for  the  tadpoles,  thus 
bringing  it  to  pass  that  if  thyroid  substance  was  specific 
organ-forming  substance,  then  frogs'  legs  were  produced  by 


144  The  Unity  of  the  Organism 

mammalian  substance,  shows  at  once  that  there  is  something 
badly  askew  in  the  theory.  So  we  are  incited  to  examine 
the  facts,  and  particularly  the  reasonings,  carefully. 

The  following  is  taken  from  one  of  Gudernatsch's  papers : 
"The  most  striking  and  at  the  same  time  unquestionable  re- 
sults were  attained  by  thyroid  feeding.  .  .  .  The  influence 
of  the  thyroid  food  was  such  that  it  stopped  any  further 
growth  but  on  the  contrary  led  to  an  abnormal  diminution 
of  the  size  in  the  animals  treated,  while  simultaneously  it  ac- 
celerated the  diff*erentiation  of  the  body  immensely  and 
brought  it  to  a  premature  end."  ^  In  other  words,  the  effect 
of  thyroid  food  was  to  stop  the  increase  in  size  of  the  frog's 
larva  and  start,  almost  at  once,  its  transformation  into  the 
adult.  Now  this  transformation  does  not  consist  merely  in 
the  production  of  legs,  but  in  a  whole  series  of  changes, 
some  of  which,  like  leg  transformation,  are  progressive, 
while  others  are  regressive.  For  example,  one  of  the  re- 
gressive changes  to  which  Gudernatsch  gives  particular 
attention  is  the  resorption  of  the  tail.  "Reduction  of  the 
body  mass  (resoi^jDtion  of  the  tail,  loss  of  water,  therefore 
an  increasing  compactness  of  the  body,  etc.)"*  more  than 
in  normal  development,  the  author  says,  goes  hand  in  hand 
with  the  progressive  changes.  That  is,  when  the  entire  series 
of  results  of  thyroid  feeding  are  considered,  and  not  merely 
one  result  picked  out  arbitrai'ily,  then  in  case  we  choose  to 
say  the  thyroid  substance  is  organ-foiTning  as  regards  legs, 
we  should  have  to  say  it  is  organ-destroying  as  regards 
tail.  Furthermore,  if  we  call  the  thyroid  substance  organ- 
fonning,  consistency  would  compel  us  to  recognize  that  one 
and  the  same  substance  is  not  only  formative  of  legs  but 
of  numerous  other  organs  and  parts,  as  of  the  skin,  mouth, 
respiratory  and  blood  systems,  all  of  which  undergo,  as  is 
well  known,  progressive  changes  during  metamoi-phosis.  Nor 
is  this  complex  series  of  morphological  changes  the  while 
story.      Striking  and  characteristic   changes   in  the  habits 


Significance  of   the  Internal  Secretory  System     145 

of  the  tadpoles  resulted.  "Towards  the  end  of  metamor- 
phosis the  animals  hardly  moved  about  in  the  water.  Tliey 
were  always  lying  quietly,  generally  on  Huir  hacks.  When 
disturbed  they  would  move  for  a  few  seconds  in  a  somewhat 
convulsive  manner  and  tlien  drop  again  to  the  bottom  of 
the  dish,  while  tadpoles  fed  on  otlier  material  would  swim 
about  for  a  long  time."  ^  So  mammalian  thyroid  substance 
is  not  only  organ-forming  for  a  whole  series  of  frog  organs 
but  it  is  habit-forming  for  a  variety  of  frog  habits ! 

But  Ave  must   not   let   the   ludicrousness    of   this    veer   us 
away   from   the   reasoning;   in    the   case.      Takin";   the    facts 

.,'0  o 

actually  brought  out  by  Gudernatsch,  what  becomes  of  the 
specificity  of  the  substance,  which  according  to  Loeb's  state- 
ments was  what  Sachs  hypothesized.^  "Sachs  suggested  that 
there  must  be  in  each  organism  as  many  specific  organ-fonn- 
ing  substances  as  there  are  organs  in  the  body."  ^'  The  trutli 
appears  to  be  that  thyroid  substance  in  tliis  case  is  organ- 
forming  in  much  the  same  sense  that  water  is  organ-forming 
for  the  leaves,  flowers,  and  fruit  in  a  squash  vine,  which  could 
not  develop  without  water.  Indeed  the  analogy  suggested 
goes  further  than  appears  at  first  sight.  As  everybody 
knows,  the  effect  on  young  plants  of  a  scant  water  supply 
is  to  stunt  the  plant  as  to  size  and  to  hasten  its  blossoming 
and  fruiting.  That  is,  an  under-supply  of  water  has  an 
effect  on  immature  plants  similar  to  that  of  an  over-su])ply 
of  thvroid  substance  on  immature  frogs,  namelv  that  of  re- 
tarding  growth  and  hastening  metam()r|)hosis.  The  total 
effect  in  each  case  is  systemic.  In  other  words,  tiie  real 
significance  of  the  instance  used  by  I.oeb  is  just  the  opjiosite 
of  his  interpretation  of  it.  Thyroid  substance  is  oryau- 
forming  only  through  being  organism-transforming.  Full 
justification  of  this  way  of  interpreting  the  ])art  in 
development  played  by  thyroid  substance  is  furnished  by  the 
recent  studies  of  15.  ^\.  Allen.  This  investigator  has,  like 
Gudernatsch,  experimented  with  frog  larva\      lie  has,  how- 


146  The  U7iity  of  the  Organism 

ever,  supplemented  Gudernatsch's  work  by  depriving  the 
larvas  of  thyroid  altogether  instead  of  giving  them  an  extra 
allowance  through  feeding.  Allen  extirpates  the  entire  em- 
bryonic thyroid  from  tadpoles  long  before  any  indications  of 
metamorphosis  appear.  What  he  finds  of  special  impor- 
tance for  the  present  discussion  is  contained  in  the  follow- 
ing; 

"While  Gudernatsch  showed  that  thyroid  feeding  acceler- 
ates development,  this  work  shows  that  the  total  absence  of 
the  thyroid  gland  produces  complete  cessation  of  somatic 
differentiation  at  a  certain  stage  but  does  not  hinder  con- 
tinued o-rowth  in  size."  ^ 

The  first  part  of  this  statement  taken  alone  might  be 
looked  upon  as  confirmatory  of  the  "formative  stuff"  theory 
of  thyroid  substance.  But  the  phrase  "at  a  certain  stage" 
implies,  as  Allen's  paper  as  a  whole  conclusively  shows,  that 
the  exact  opposite  of  such  a  conception  is  alone  tenable. 
What  actually  happens,  Allen  brings  to  light,  is  that  in 
spite  of  the  complete  absence  of  thyroid  substance  trans- 
formation of  the  larva  is  begun  but  is  not  carried  through. 
That  is,  the  organs  and  parts  of  the  adult  frog  are  laid 
down  but  (with  the  exception  of  the  reproductive  glands) 
are  not  completed.  "It  is  evident,"  Allen  says,  "that  the 
thyroid  gland  is  in  no  wise  essential  to  the  earlier  phases  of 
development,  but  that  at  a  certain  definite  stage,  further 
development  of  the  soma  is  dependent  upon  it."  '^  It  should 
be  mentioned  that  Allen  makes  a  rather  special  point  of  the 
accordance  of  his  results  with  those  of  Gudernatsch. 

If  the  greatly  hastened  and  modified  metamorphosis  of  the 
frog  tadpole  observed  by  Gudernatsch  is  inducible  by  no 
other  means  than  by  thyroid  substance  (which  while  pos- 
sible is  not  at  all  certain),  then  is  the  substance  causal  in 
the  important  sense  of  being  not  only  competent  but  in- 
dispensable. But  even  so  it  falls  far  short  of  being  a  com- 
plete causal  explanation  of  the  phenomena  under  contempla- 


Significance  of   the  Internal  Secretory  System     147 

tion,  as  Allen's  results  directly  prove. 

A  defect  in  Loeb's  reasoning  is  liis  ignoring  the  truth  that 
thyroid  substance  lacks  in  the  case  cited  tlie  prime  attri})ute 
of  a  sufficient  cause,  namely  full  competency.  The  actual 
substance  which  enters  into  the  new  legs,  and  as  far  as  that 
goes,  into  the  other  new  parts,  is  probably  provided  very 
little,  if  at  all,  by  the  specific  substance  or  hormone  of  the 
thyroid.  The  causal  role  played  is  the  relatively  humble  one 
of  excitor  or  stimulator  to  an  activity  not  essentially  new 
but  only  exceptional  as  to  time.  Of  course  Loeb  does  not 
need  to  be  told  that  hormones  incite  growth  rather  than 
provide  the  substance  itself  out  of  which  the  organs  and 
parts  are  built.  Indeed  portions  of  his  account  of  this  very 
case  show  positively  that  he  is  aware  of  this.  We  read : 
"Thus  we  see  that  the  mesenchyme  cells  giving  rise  to  legs 
may  lie  dormant  for  months  or  a  year  but  will  grow  out 
when  a  certain  type  of  substances,  e.g.,  thyroid,  circulates  in 
the  blood.  There  may  exist  an  analogy  between  the  activ- 
ating effect  of  the  thyroid  substance  and  the  activating 
effect  of  the  spermatozoon  or  butyric  acid  (or  other  par- 
thenogenetic  agencies)  upon  the  Qgg.'^'' ^ 

This  suggestion  of  analogy  between  the  action  of  thyroid 
secretion  and  "butyric  acid  (or  other  parthcnogenetic  agen- 
cies)" is  well  taken.  The  resemblance  between  the  two 
agencies,  as  judged  by  their  effects  on  development,  is  cer- 
tainly rather  close.  Very  well ;  would  Loeb,  then,  call  bu- 
tyric acid  organ-forming  substance,  and  identify  it  with  the 
"formative  stuff"  of  Sachs  .'^  Certainly  any  substance  which 
will  rouse  the  latent  developmental  cai)acitics  of  an  Qgg  into 
activity  is  in  a  minor  sense  formative,  especially  if  these 
capacities  are  wholly  unable  to  start  without  some  such 
agency.  But  since  butyric  acid,  or  some  one  of  tlie  other 
dozens  or  scores  of  parthcnogenetic  agencies,  may  activate 
the  eggs  of  many,  many  species  of  animals;  and  since  the 
eggs  of  many,  many  species  may  be  ai^tivated  by  any  one  of 


148  The  Unity  of  the  Organism 

scores  of  such  agencies,  what  about  the  specificity  of  the 
formative  substances  which  Loeb  himself  expressly  says  was 

A^rt  of  Sachs'  conceptions? 
jj^gain,   how   reconcile   the    contention    that   thyroid   sub- 

^^ts^e  or  something  like  it  is  organ-forming  for  the  legs 
olM-  ^SWpole  witli  the  statement  quoted  a  few  paragraphs 
bac^aljjlit  mesenchyme  cells  which  give  rise  to  legs? 

The  ^Md^jmy steal  and  Logical  Weakness  of  the  View 

Obviously  Loeb's  treatment  of  this  subject  contains  irre- 
conciliable  contradictions.  Is  it  then  worthless?  My  answer 
is,  no,  not  by  any  means.  But  how  comes  it  that  a  scien- 
tist of  his  great  experience  and  merited  distinction  can  run 
into  such  self-destroying  speculations  and  statements,  seem- 
ingly without  rational  discomfort  to  either  himself  or  others 
of  the  school  which  he  represents? 

The  answer  takes  us  back  to  some  of  the  most  funda- 
mental issues  between  the  elementalist  and  organismal  stand- 
points, and  though  not  requiring  us  to  palliate  in  any  degree 
such  offenses  against  scientific  reasoning,  it  partly  explains 
how  tolerance  for  such  oflTenses  is  begotten,  and  discovers 
a  nucleus  of  genuine  merit  in  Loeb's  position.  Bringing  the 
matter  to  as  basal  a  statement  as  possible,  what  we  find  is 
that  this  whole  book  on  the  Organism  as  a  Whole  is  written 
on  the  theory  that  the  only  alternative  to  the  assumption 
of  supernaturalism  is  materialism.  Instead  of  supernatural 
forces  of  some  sort  (Platonic  Ideas,  entelechies,  psychoids, 
"supergenes")  for  explaining  the  organism  when  regarded 
as  alive  and  whole,  his  assumption  is  that  material  elements 
as  known  to  us  in  inorganic  nature  are  the  sufl[icient  causal 
explanation  of  organic  phenomena. 

Were  this  theory  correct — were  it  true  that  the  "vital 
principle"  must  either  be  conceived  as  supernatural  or  tliat 
the  inorganic  elements  taken  by  themselves   are  competent 


Significance  of   the  Internal  Secretory  Sij stent      Wd 

to  produce  organisms,  then  it  would  be  impossible  for  biol- 
ogy to  do  much  better  in  its  reasoning  and  general  attitude 
than  Loeb  and  other  elementalists  do  when  they  undertake 
to  construct  a  philosophy  of  organisms.  I  agree  wliole- 
heartedly  that  all  supernaturalism,  no  matter  what  nomen- 
clatorial  garb  it  takes  on,  must  be  repudiated  by  the  sciences 
of  organic  beings.  Ideas,  or  psyclioids,  or  entelechies,  or 
"principles"  of  any  kind  conceived  as  independent  of,  or 
even  separable  from,  sensible  objects  are  quite  as  repugnant 
to  me,  an  organismalist,  as  they  are  to  any  elementalist.  The 
essence  of  my  contention  is  that  the  natural  substitute  for 
these  imponderable  things  are  the  living,  individual  organ- 
isms themselves,  and  not  the  particles  of  which  they  are  com- 
posed. Each  and  every  individual  organism  is  a  natural 
reality  by  exactly  the  same  criteria  that  the  atoms,  mole- 
cules, cells  and  tissues  of  which  it  is  composed  are  natural 
realities.  And  since  each  individual  is  to  some  extent  differ- 
ent from  every  other,  and  maintains  its  individuality  in  full 
possession  of  these  differences,  by  its  power  of  transforming 
foreign  substance  into  its  own  substance,  it  is  ultimate  both 
as  to  structure  and  as  to  causal  power  in  as  deep  and  literal 
a  sense  as  the  material  particles  of  which  it  is  composed  are 
ultimate. 

Loeb's  considerable  attention  to  the  views  of  Claude  Ber- 
nard is  fortunate  for  us,  since  it  affords  a  chance  to  show 
from  still  another  angle  the  inevitable  breakdown  of  ele- 
mentalist reasoning  when  it  is  brought  face  to  face  with  or- 
ganic phenomena  as  actual  nature  presents  them  to  tiie 
modern  student.  Loeb  calls  attention,  pro})erly,  to  the  fact 
that  one  of  the  things  on  which  Bernard  ]ihu'ed  special  em- 
phasis, as  Bichat  before  him  had  done,  is  the  organizing  syn- 
theses which  go  on  in  the  living  being.  The  real  advance,  it 
seems  to  me,  which  Bernard  made  over  any  of  his  ])re{leces- 
sors,  was  the  positiveness  of  his  rejection  of  a  vital  force 
as    something    supernatural— as    something,    using   hi-;    own 


150  The  Unity  of  the  Organism 

words,  "under  the  government  of  a  special  principle,  a  pecu- 
liar power  what  name  soever  be  given  it,  whether  soul,  or 
archeon,  or  psyche,  or  plastic  intermediary,  or  guiding 
spirit,  or  vital  force,  or  vital  properties";^  and  his  rejec- 
tion of  these  conceptions  because  of  his  recognizing  that  the 
unitariness  of  the  organism  removes  the  necessity  for  as- 
suming any  such  extraneous  principle.  "When  we  say," 
Bernard  writes,  "that  life  is  a  guiding  idea,  or  the  evolu- 
tive force  of  the  being,  we  merely  express  the  thought  of  a 
unity  in  the  succession  of  all  the  morphological  and  chemical 
changes  effected  by  the  germ,  from  the  beginning  to  the 
end  of  life.  Our  mind  grasps  that  unity  as  a  conception  it 
cannot  escape.   .   .   ."  ^^ 

Up  to  this  point  the  position  held  by  Bernard  is  entirely 
satisfactory  for  the  organismal  conception  as  I  am  trying 
to  develop  it,  and  is,  according  to  my  view,  unassailable. 
But  the  unquoted  part  of  the  last  sentence  contains  a  state- 
ment which  reveals  Bernard  on  a  by-road  leading  away  from 
the  promised  land  toward  which  he  w^as  headed  as  long  as  he 
was  speaking  in  terms  of  biology  proper.  The  rest  of  tlie 
sentence  follows :  "and  explains  it  as  'a  force' ;  but  the  mis- 
take is  in  supposing  that  this  metaphysical  force  acts  after 
the  manner  of  a  physiological  force."  ^^  Stated  in  a  nut- 
shell, the  by-road  which  Bernard  is  entering  here  is  that  of  a 
kind  of  separatedness,  but  inevitable  concomitance,  or  paral- 
lelism between  the  phenomenal  and  neumonal  worlds  which, 
according  to  the  views  upheld  in  this  volume,  does  not  exist. 
"We  need  here,"  Bernard  says,  "to  draw  a  distinction  be- 
tween the  metaphysical  world  and  the  phenomenal  physical 
world,  which  serves  as  its  basis,  but  which  can  borrow  noth- 
ing from  it."  This  Leibnitzian  theory,  according  to  which 
"everything  takes  place  in  the  soul  as  though  there  were  no 
body,  and  in  the  body  everything  takes  place  as  thougli 
there  were  no  soul,"  Bernard  says  science  "recognizes  and 
adopts  in  our  day."     But  in  our  day,  this  year  nineteen 


Significance  of   the  Internal  Secretory  System     151 

hundred  and  eighteen,  science,  at  least  so  much  of  it  as 
speaks  through  tliis  vohnne,  tliou^li  understanding  fully 
what  this  dualist  theory  is,  rejects  it.  It  denies  that  things 
take  place  in  the  soul  as  though  tlure  were  no  hody;  and 
that  things  take  place  in  the  body  as  though  tliere  were  no 
soul.  On  the  contrary  it  affirms  that  what  in  rather  un- 
critical language  we  call  "the  hody"  and  "tlie  soul"  are  in 
the  most  intimate  and  indissoluble  connection  with  and  de- 
pendence upon  each  other,  and  together  constitute  "the 
Orcranism." 

o 

The  Form  of  Metaphysical  Absolutism  Involved 

A  full,  systematic  justification  of  this  position  is  beyond 
the  province  of  this  volume;  and  in  this  chapter  we  are  con- 
cerned solely  with  the  "body,"  the  strictly  morphological 
and  physiological  aspect  of  the  subject.  Nevertheless,  this 
much  of  contact  with  the  "soul"  aspect  was  unavoidable  for 
the  reason  that  Bernard  and  also  Locb  have  run  into  it  in 
such  fashion  as  to  color  deeply  their  discussions  and  out- 
look. This  coloring  is  the  more  unfortunate  and  the  more 
insistent  in  requiring  attention  from  the  fact  tliat  the  au- 
thors, especially  Loeb,  are  apparently  unaware  of  such 
coloring.  For  example,  Loeb  writes  on  the  first  page  of  his 
book,  after  saying  that  the  atomistic  theory  of  matter  and 
electricity  are  now  in  all  probability  on  a  "permanent 
basis":  "This  permits  us  to  state  as  an  ultimate  aim  of  the 
physical  sciences  the  visualization  of  all  phenomena  in  tenns 
of  groupings  and  displacements  of  ultimate  particles,  and 
since  there  is  no  discontinuity  between  the  matter  consti- 
tuting the  living  and  non-living  world  the  goal  of  biology 
can  be  expressed  in  the  same  way." 

Statements  like  this,  many  of  which  can  easily  he  ([uoled 
from  Loeb's  writings,  leave  no  question  about  his  meta- 
physical affinities.     The  conception  of  "ultimate  particles" 


152  The  Unity  of  the  Organism 

as  explanation  of  all  phenomena,  is  exactly  what  I  mean  by 
elementalist  absolutism. 


Confusion  of  Theory  of  Organisms  and  Theory  of  the 

Knowledge  of  Organisms 

Although  the  thoroughgoing  metaphysical  character  of 
these  statements  is  evidenced  by  the  finalism  which  crops 
out  at  several  points,  this  is  not  the  aspect  of  the  matter 
which  chiefly  interests  us  here.  Rather  what  we  are  con- 
cerned with  is  the  fact  that  affirmations  about  the  "aim  of 
the  physical  sciences"  and  the  "goal  of  biology"  do  not 
belong,  properly  speaking,  to  the  provinces  of  physical 
science  and  biology  at  all,  but  to  quite  a  different  science, 
namely  that  which  deals  with  the  nature  of  knowledge  itself. 
The  "physical  sciences"  are  the  vast  accumulation  of  man's 
positive  knowledge,  theories,  hypotheses,  and  so  forth,  about 
physical  nature;  they  certainly  are  not  physical  nature  it- 
self. Consequently  a  statement  of  the  character  and  aims 
of  that  knowledge  is  not  a  statement  about  the  phenomena 
to  which  the  knowledge  pertains.  And  the  same  reasoning 
applies  to  the  affirmation  about  the  goal  of  biology. 

All  this  is  only  another  way  of  saying  what  Loeb  him- 
self virtually  tells  us,  namely  that  his  entire  discussion  of 
the  organism  as  a  whole  is  made  from  the  standpoint  of  one 
particular  theory  of  the  ultimate  nature  of  living  beings, 
that  theory  being  the  mechanistic.  Recall  the  complete  title 
of  the  book  The  Organism  as  a  Whole  From  fhe  Physico- 
Chemical  Viewpoint.  To  treat  the  subject  from  this  view- 
point is  of  course  perfectly  legitimate.  When,  however,  the 
assumption  is  made  that  such  a  treatment  is  the  only  really 
legitimate  one  because  it  rests  on  idtimate  truth,  then  sound 
science  is  bound  to  protest,  chiefly  because  of  the  obvious 
biological  inadequacy,  and  at  some  points,  perversion  and 
contradiction  displayed  in  the  treatment. 


Significcmce  of   the  Internal  Secretory  System     153 

This  brings  us  back  to  the  way  internal  secretions  are 
dealt  with  bj  Loeb.  His  failure  to  (Hstlnguisli  between  tlie 
two  very  distinct  fields  of  theory  above  indicated,  namely 
theories  about  the  phenomena  of  living  beings,  and  theories 
about  knowledge  of  these  piienomcna,  largely  explains  the 
defects  of  the  theoretical  parts  of  his  work.  And  we  are 
now  in  position  to  give  our  criticism  greater  expllcitness. 
Consider  for  example  in  tlie  light  of  what  has  just  been  said 
about  confusion  relative  to  kinds  of  theory,  the  irreconcil- 
able statements,  cited  on  an  earlier  page,  that  the  mesen- 
chyme cells  give  rise  to  the  legs  of  the  tadpole,  and  that 
thyroid  substance  is  organ-forming  substance  for  the  legs. 
Stated  briefly  the  case  seems  to  be  thus :  Loeb  recognizes,  as 
every  one  must,  that  internal  secretions  constitute  a  physico- 
chemical  agency  for  bringing  about  that  harmonious  de- 
velopment and  functioning  so  characteristic  of  the  organ- 
ism. But  this  harmony  is  one  of  the  very  things  which  has 
seemed  to  some  biologists  inexplicable  w^thcxit  the  assum]>- 
tion  of  supernatural  influences  of  some  sort;  hence  Driesch's 
attempt  to  modernize  the  ancient  entelechy.  But  since  in- 
ternal secretions  play  the  role  that  entelechies  are  sup})osed 
to  play,  namely  that  of  establishing  and  maintaining  the 
unity  and  equilibrium  of  the  organism,  the  need  for  ente- 
lechies no  longer  exists ;  at  least,  this  would  be  so  for  all 
persons  who  do  not  contend  that  "ultimate  explanation"  is 
the  "goal  of  science." 

To  those  who  hold  these  absolutist  beliefs  as  to  the  power 
and  aims  of  science  three  favorite  courses  are  open  and  are 
followed  by  different  representatives  of  the  school,  depend- 
ing on  the  taste,  training  and  outlook  of  the  person.  One 
course  consists  in  pointing  out,  taking  an  illustrative  case, 
that  internal  secretions,  being  only  contributing  causes,  do 
not  constitute  an  ultimate  ex])lanation,  so  tliat  entelechies 
or  something  similar  arc  as  necessary  as  l)i'f()rc.  'I'his  would 
be  the  course  followed  by  the  vitalistic  wing  of  the  ahsolu- 


154  The  Unity  of  the  Orgcmism 

tist  school.  For  them  internal  secretions  would  be,  in  so  far 
as  they  contribute  to  the  harmony  of  the  organism,  merely 
agencies  produced  and  used  by  supernatural  causes. 

Another  course,  and  perhaps  the  one  most  frequented  by 
elementalists,  would  be  to  contend  that  internal  secretions 
are  sufficient  as  a  causal  explanation  of  organic  unity  to 
make  the  entelechy  or  any  similar  notion  quite  superfluous, 
even  though  these  substances  are  far  from  a  complete  ex- 
planation. The  reasoning  of  this  group  of  elementalists  as 
to  this  situation  is  substantially  as  follows :  although  in- 
ternal secretions  fall  far  short  of  fully  explaining  organic 
unity  and  harmony,  the  action  of  these  being  merely  that 
of  incitors  and  inhibitors,  they  are  yet  genuinely  causal, 
genuinely  physico-chemical  and  so  are  on  the  road  toward 
complete  explanation  of  the  phenomena.  All  that  is  nec- 
essary consequently,  is  to  believe  that  still  further  advance 
in  the  same  direction  will  reach  finallv  a  full  elementalistic 
explanation ;  that  is,  an  explanation  which  will  have  no  need 
of  either  supernatural  elements  or  the  organism  as  a  whole. 
The  attitude  of  this  large  class  is  one  primarily  of  faith 
rather  than  of  reliance  on  positive  knowledge ;  they  are 
inspired  more  by  what  they  believe  they  will  do  in  the  future 
than  by  what  they  actually  have  done.  They  are  preemi- 
nently men  of  promises.  Although  their  achievements  in 
experimental  science  are  indeed  large,  the  results  reached 
by  them  are  prized  more  on  account  of  what  they  are  believed 
to  augur  for  the  future  than  for  their  present  meaning  and 
worth. 

Then,  finally,  there  is  the  group  of  elementalistic  absolu- 
tists, of  whom  the  author  of  The  Organism  as  a  Whole  from 
the  Physic o-Chemical  Viewpoint  is  one  of  the  most  eminent 
in  our  day,  w^ho,  as  we  have  pointed  out,  by  confusing  theo- 
ries about  objective  phenomena  witli  theories  about  the 
knowledge  of  such  phenomena,  are  led  to  affirm  that  such 
phenomena  as  the  unity  of  the  organism  are  fully  explained 


Significance  of   the  Internal  Secretory  System     155 

by  internal  secretions,  Avhcn  as  a  matter  of  fact  tliey  are 
only  partly  explained  thus. 

The  surprising  tiling  about  the  confusion  of  this  group 
is  that  wholly  irreconcilable  positions  are  held  with  ini[)uii- 
ity,  such  for  instance  as  those  according  to  which  thyroid 
substance  is  held  to  be  the  organ-forming  substance  of  frogs' 
legs  in  one  part  of  a  discussion,  and  mesenchyme  cells  are 
acknowledged  to  be  of  this  nature  in  another  part.  Tlic 
contradiction  is,  to  be  sure,  often  of  such  character  as 
easily  to  escape  the  uncritical  reader;  but  as  to  the  au- 
thors of  such  contradictions  no  other  explanation  seems 
possible  than  that  of  wrong  habits  of  scientific  thought  be- 
gotten of  untenable  a  priori  conceptions.  For  example,  a 
hasty  reading  of  the  discussion  under  review  might  lead  one 
to  suppose  that  thyroid  substance  is  not,  after  all,  regarded 
by  Loeb  as  anything  more  than  one  contributing  cause  of 
frogs'  legs,  mesenchyme  cells  being  recognized  as  another 
cause.  Close  attention  to  the  text  does  not,  however,  war- 
rant this  generous  interpretation  of  the  author's  position. 
Going  back  to  his  espousal  of  the  theory  of  Sachs  and  other 
botanists  as  to  organ-forming  substances,  we  read :  "Specific 
shoot-producing  substances  are  carried  to  the  apex,  while 
specific  root-producing  substances  are  carried  to  the  base  of 
a  plant.  When  a  piece  is  cut  from  a  branch  of  willow  the 
root-forming:  substances  must  continue  to  flow  to  the  basal 
end  of  the  piece,  and  since  tlieir  further  progress  is  blocked 
there  they  induce  the  formation  of  roots  at  the  basal 
end."  1^ 

If  this  means  anything  it  means  that  the  shoots  and  roots 
are  actually  built  up  by  material  carried  about  in  the  wil- 
low branch.  There  is  nothing  in  the  language  tliat  can  he 
interpreted  as  meaning  that  shoot-and-root-forming  sub- 
stances are  mere  stimulators  of  some  other  substances  which 
become  the  actual  shoots  and  roots.  Yet  it  is  witli  forma- 
tive  substances   of   this    sort   that  Loeb  in   some   parts    of 


156  The  Unity  of  the  Organism 

his  discussion  explicitly  identifies  internal  secretions.  "At 
that  time  the  idea  of  the  existence  of  such  specific  organ- 
forming  substances  was  received  with  some  skepticism.  .  .  . 
Such  substances  are  known  now  under  the  name  'internal 
secretions'  or  'hormones' ;  their  connection  with  the  theory 
of  Sachs  was  forgotten  with  new  nomenclature."  ^  Then 
follows  the  reference  to  the  tadpole  legs ;  so  that  were  consis- 
tency really  a  jewel  to  the  author,  he  could  not  escape  mean- 
ing that  the  substance  of  the  legs  was  actually  derived  di- 
rectly from  the  particular  internal  secretion  in  question,  in 
this  case  thyroid  substance;  and  the  later  statement  about 
the  formation  of  legs  from  dormant  mesenchyme  cells 
through  the  mere  activating  effect  of  thyroid  substance  is  by 
implication  contradicted. 

This  brings  up  again  a  matter  about  the  interpretation 
of  development  which  we  have  dwelt  upon  in  several  other 
connections,  that  of  protest  against  rejecting  the  indubi- 
table evidence  of  the  senses  in  favor  of  a  priori  conceptions. 
The  crucial  question  in  the  present  case  is  this :  which  is  the 
more  fundamentally  organ-forming  substance  for  frogs'  legs, 
the  mesenchyme  cells  which  "though  giving  rise  to  legs  may 
lie  dormant  for  months,"  or  the  thyroid  substance  which 
may  stimulate  these  cells  into  premature  activity?  While 
Loeb's  discussion  does  not  raise  this  question  definitely,  the 
implication  is  unescapable  that  thyroid  substance  is  for  him 
the  more  fundamental.  What  else  is  the  meaning  of  the 
contention  that  this  substance  is  organ-forming  while  no- 
where do  we  find  the  mesenchyme  cells  so  designated?  Yet 
the  observational  evidence  is  that  the  production  of  legs  is 
accomplished  through  the  transformation  of  mesenchyme 
and  other  cells  which  in  the  larva  are  not  leg-substance,  but 
in  the  adult  are  leg-substance.  Hence  it  follows  that  so  far 
as  actual  observation  is  concerned  the  mesench^^matous  and 
other  larval  substances  are  more  entitled  to  be  called  organ- 
forming  than  is  the  thyroid  substance. 


Significance  of   the  Internal  Secretory  System     157 

An  Illustration  of  Neglect  of  Fact  By  ElevientaUst  Theory 

Tliesc  reflections  lead  to  a  still  deeper  level  of  the  inherent 
faultiness  of  elemcntalistic  absolutism  in  biolo'^y,  the  toler- 
ance which  it  engenders  for  ignoring  relevant  facts;  or 
stated  otherwise,  of  arbitrarily  selecting  from  a  great  com- 
plex of  facts  just  those  which  suit  the  argument,  and  dis- 
regarding all  the  others.  For  example,  recall  Loeb's  ref- 
erence to  precocious  leg-production  in  frog  tadpoles  as 
though  the  effect  of  thyroid  feeding  stood  alone  rather  than 
as  one  among  a  great  concatenation  of  effects,  some  con- 
structive and  some  destructive,  this  complex  of  phenomena 
constituting  the  metamorphosis  of  the  young  into  the  adult. 
Loeb's  use  of  Gudernatsch's  results  amounts  to  a  positive 
obscuration  for  the  reader  of  what  these  important  experi- 
ments really  teach.  Only  by  the  culling  of  facts  to  suit  the 
argument  and  the  use  of  certain  words  and  phrases,  as 
"influence,"  "responsible  for,"  and  so  on,  with  equivocal 
meanings,  can  these  results  be  made  to  support  the  conten- 
tion that  thyroid  substance  is  specific  organ-forming  sub- 
stance for  frogs'  legs.  The  patent  fact  is  that  certain 
mesenchyme  and  other  cells  of  the  larva  are  organ-forming 
for  legs,  and  there  is  no  straightforward  way  of  talking 
about  the  causes  of  the  transformation  of  a  given  grou])  of 
more  or  less  undifferentiated  tadpole  cells  into  the  much 
enlarged  and  highly  differentiated  group  called  a  leg,  with- 
out recognizing  the  whole  organism  as  causal  of  the  ])ar- 
ticular  transformation.  Probably  no  set  of  discoveries  con- 
cerning the  development  of  the  individual  has  ever  Ix^en 
made  which  so  objectifies  the  means  employed  by  the  whole 
in  producing  and  correlating  its  constituent  parts  as  those 
on  internal  secretions ;  and  not  the  least  significant  fact  is 
that  these  substances  are  themselves  produced  by  tlie  or- 
ganism. 

Even  were  Loeb's  contention  valid  that  thvroid  substance 


158  The  Unit?/  of  the  Organism 

is  specific  organ-forming  substance,  the  indubitable  fact  that 
in  normal  development  tliis  substance  is  itself  a  product  of 
the  organism's  activities,  throws  it  into  a  very  subordinate 
place  as  a  cause  of  development.  Tlie  truth  is,  the  main 
upshot  of  the  effort  to  explain  ontogeny  on  elementalistic 
principles  amounts  to  an  effort  to  avoid  recognizing  the  most 
positive  and  definite  entity  in  the  whole  situation,  namely 
the  organism  taken  alive,  normal  and  untampered  with. 

A   Peculiar  ElementaUst   Objection  to   the  Organic    Whole 

We  now  come  to  the  last  point  to  be  noticed  in  connec- 
tion with  the  elementalist  attempt  to  deal  with  internal  se- 
cretions as  related  to  the  organism  as  a  whole.  Instancing 
the  familiar  way  in  which  a  particular  part  of  a  flat-worm 
will  give  rise  to  a  new  head  after  being  cut  away  from  tlie 
original  animal,  when  no  head  would  have  been  formed  at  this 
place  had  not  the  animal  been  cut,  Loeb  writes :  "How  does 
the  'whole'  suppress  all  this  formative  power  in  the  part  be- 
fore the  latter  is  isolated?  It  almost  seems  as  if  the  isola- 
tion itself  were  the  emancipation  of  the  part  from  the  tyr- 
anny of  the  whole.  The  explanation  of  this  tyranny  or  of 
the  correlation  of  the  parts  in  the  whole  is  to  be  found,  how- 
ever, in  a  different  influence."  ^^ 

Then  follows  the  statement  previously  quoted  about  the 
specific  organ-forming  substances  of  Sachs  and  other  bot- 
anists, and  the  assumed  identity  of  these  with  internal  se- 
cretions. 

Without  raising  the  question  concerning  the  evidence  for 
the  assumption  that  the  production  of  a  flat-worm's  head 
as  indicated  is  dependent  upon  internal  secretions,  let  us 
consider  a  moment  the  interesting  conception  thrown  into 
the  treatment  that  the  whole  flat-worm  tyrannizes  over  its 
parts.  Why  this.'^  Is  it  "mere  rhetoric".'^  We  are  not  per- 
mitted to  judge  it  thus,  for  no  one  has  pronounced  against 


Significance  of   the  Internal  Secretory  System     159 

this  sort  of  thing  in  science  more  frequently  than  Loeb.  The 
conception  that  the  "whole"  exercises  a  "tyranny"  over  the 
parts  we  must  accept  as  being  seriously  scientific  with  Loeb. 
Well  then,  since  tyranny  is  "absolute  power  arbitrarily  or 
unjustl}^  administered,"  according  to  the  dictionary,  it  is 
certainly  interesting  to  an  organismalist  to  find  so  eminent 
an  elementalist  acknowledging  the  organism  as  a  whole  to  be 
truly  causal  relative  to  its  parts — for  it  is  hardly  conceiv- 
able that  even  the  extreme  pliability  of  elementalist  practice 
as  regards  the  definition  of  words  would  venture  to  hold  the 
absolute  power  which  constitutes  tyranny  to  be  without 
causal  efficiency.  Power  thus  potent  but  which  could  cause 
nothing,  not  even  the  destruction  of  the  parts  (for  surely 
the  tyranny  of  the  whole  does  not  destroy  the  parts),  would 
be  too  queer  a  conception  for  anybody  to  father  deliber- 
ately. 

But  the  most  interesting  thing  about  this  idea  of  the 
"tyranny"  of  the  whole  over  the  parts  remains  to  be  no- 
ticed. T3^ranny  is  not  merely  absolute  power;  it  is  such 
power  exercised  unjustly  or  "in  a  manner  contrary  to  law 
or  justice." 

Here  we  come,  I  think,  to  about  the  last  ditch  of  the  ele- 
mentalist position.  On  what  ground  does  one  conceive  the 
power  exercised  by  the  whole  organism  over  its  parts,  to  be 
contrary  to  law  or  justice.  According  to  what  legisla- 
ture or  court  is  there  a  law  of  the  parts  of  an  organism  more 
just  than  the  law  of  the  whole?  None  whatever  in  nature, 
it  must  I  think,  be  admitted.  The  only  ground  for  the  ele- 
mentalist's  pronouncement,  that  the  wliole  acts  tyrannously 
toward  the  parts,  that  it  acts  in  a  manner  "contrary  to  law 
or  justice"  is  in  the  mind  of  the  yerson  who  makes  the  pro- 
nouncement. 

The  truth  is — and  it  is  of  great  importance  since  its 
influence  reaches  far  beyond  the  confines  of  scientific  tech- 
nicalities— any    scientist,    especially    any    biologist,    who    is 


160 


The  Unity  of  the  Organism 


through  and  through  an  elementalist,  is  necessarily  a  protes- 
tant  against  all  law  except  the  law  of  elements. 

The  scientific  elementalist  is  inevitably  anarchistic  toward 
all  the  most  common,  most  objective,  structures  and  laws  of 
nature. 

His  faith  is  in  the  laws  of  the  obscure  or  invisible  world 
and  against  those  of  the  everywhere  visible  world.  Atoms 
are  more  real  to  his  mind  than  are  lands  and  waters,  plants 
and  animals. 

REFERENCE  INDEX 


1.  Loeb   ('16)    155 

2.  Loeb   ('16)    156 

3.  Gudernatsch     475 

4.  Gudernatsch     476 

5.  Gudernatsch     476 

6.  Loeb   ('16)    159 


7.  Allen    ('18)    515 

8.  Loeb   ('16)    156 

9.  Bernard     516 

10.  Bernard     519 

11.  Loeb   ('16)    154 

12.  Loeb    ('16)     154 


Apjirvh 


AU 


jKorri 


Chapter  XX 

NEURAL  INTEGRATION 

Neural  and  Not  Fsycliical  Phenomena  the  Subject  of  ThU 

Chapter 

THE  fact  should  be  firmly  fixed  in  mind  at  the  outset 
that  in  this  chapter  we  have  nothing  to  do  with  tlic 
organism's  consciousness  and  volition.  We  are  to  deal  with 
the  nervous  system  purely  on  the  basis  of  its  physical  activi- 
ties. Whether  or  not  consciousness  or  something  of  the  es- 
sential nature  of  it  appertains  to  part  or  all  of  the  activities 
to  be  treated  we  do  not  have  to  decide,  so  far  as  this  discus- 
sion is  concerned.  Our  task  here  is  to  be  strictly  anatomical 
and  physiological,  and  not  at  all  psychological. 

Distinction  Between  Developmental  and  Functional 

Integration 

The  discussion  may  be  opened  by  calling  attention  again 
to  a  matter  noticed  incidentally  in  the  last  chapter,  namely 
the  relative  parts  played  by  internal  secretions  and  the  ner- 
vous mechanism  in  integrating  the  developing  organism  on 
the  one  hand,  and  the  functioning  organism  on  the  otiier. 
We  saw  that  the  role  of  internal  secretions  in  preserving 
developmental  equilibrium  in  the  individual  is  so  conspicu- 
ous as  to  throw  the  strictly  ])hysio-logical  role  of  tlie  sub- 
stances somewhat  into  the  background.  Nevertheless  as  evi- 
dence, particularly  clinical  evidence,  lias  made  clear,  tlieir 
part  in  functional  equilibrium  is  far-reaching. 

101 


162  The  Unity  of  the  Organism 

We  must  now  point  out  specifically  what  appeared  only 
incidentally  in  the  previous  discussion,  namely,  that  the 
integrative  action  of  hormones  and  of  the  nervous  system 
are  to  a  considerable  extent  the  reverse  of  each  other  as 
regards  their  relation  to  development  and  to  adult  func- 
tion. That  is,  while  harmones  are,  perhaps,  more  impor- 
tant in  development  and  become  relatively  less  significant 
with  the  advancing  age  of  the  individual,  the  nei-vous  sys- 
tem plays  a  minor  part  in  integrating  the  developmental 
processes  but  becomes  of  supreme  importance  in  this  way 
for  the  functioning  of  the  adult.  Or,  stating  the  generali- 
zation in  another  way,  integration  of  the  organism  is  ac- 
complished by  chemical  means  chiefly,  and  by  neural  means 
little  or  not  at  all,  during  early  life,  and  by  neural  means 
chiefly  and  by  chemical  means  secondarily  in  later  life. 

One  can  hardly  fail  to  see,  in  a  general  way,  the  bearing 
of  this  on  the  familiar  truth  that  the  life  of  the  individual 
among  the  higher  animals,  man  especially,  is  successively 
vegetal,  animal,  emotional,  and  finally  rational  and  intel- 
lectual in  its  dominant  characteristics,  with  the  successive 
stages  of  earlier  and  later  childhood,  youth,  and  earlier 
and  later  maturity. 

The  Author's  Indebtedness  to  Sherrington's  Work 

My  dependence  upon  Sherrington's  work  in  this  field  will 
be  so  great  as  to  make  it  impossible  to  acknowledge  it  at 
every  point.  I  therefore  make  at  the  outset  the  general 
statement  that  a  large  part  of  my  discussion  consists  of  a 
re-wording  and  rearranging  of  facts  and  ideas  contained  in 
this  physiologist's  important  book,  The  Integrative  Action 
of  the  Nervous  System.  But  while  my  chief  reliance  here 
is  on  Sherrington's  work,  the  writings  of  Cannon  and  his 
collaborators  have  been  the  most  important  source  of  what 
I  have  to  say  about  tlie  autonomic  nervous  system.     Can- 


\ 


Neural  Integration  103 

non's   recent  book,  Bodily  Changes  in  Pain,   Hunger,  Fear 
and  Rage,  has  been  especially  drawn  upon. 

The  Fund  amen  tality  of  Cellidar  Integration  in  the  Reflex 

Arc 

In  no  department  of  pliysiolog'y  do  cells  keep  themselves 
more  persistentl}'^  in  the  attention  of  the  student  than  in  tlie 
physiology  of  the  nervous  system.  But  likewise  nowhere 
is  the  fundamental  dependence  of  cells  on  other  cells  more 
clearly  seen ;  for  -while,  speaking  from  the  standpoint  of 
general  functions,  cells  may  be  looked  upon  as  individual 
units,  when  it  comes  to  the  study  of  nerve  cells  as  such, 
that  is,  as  constituents  of  the  functioning  nervous  system, 
the  individual  cell  is  found  to  be  no  longer  the  basal  unit. 
Viewed  thus  the  reflex-arc  and  not  the  individual  cell  is  the 
unit. 

The  distinction  thus  indicated  is  important  from  the  or- 
ganismal  standpoint  and  must  be  considered  a  little  more 
fully.  The  general  functions  of  cells  to  which  reference  i& 
made  above  are  those  common  to  all  cells,  even  those  of  the 
tissues  of  fully  differentiated  multicellular  animals.  No 
matter  what  tissue  be  under  consideration,  muscle,  gland, 
epithelial  or  what  not,  so  long  as  it  is  truly  living,  each  and 
every  cell  assimilates,  breathes,  excretes,  and  carries  on  all 
the  metabolic  processes.  Thus  far  each  cell  is  an  indepen- 
dent unit  in  a  high  degree.  Now  while  all  tissue  cells,  using 
the  term  tissue  in  its  common  histological  sense,  have  a  rela- 
tional or  integrational  function  in  addition  to  these  indi- 
vidual functions,  it  is  in  the  nervous  system  that  this  inte- 
grative aspect  of  cell  life  is  most  positive  and  definable. 
The  reflex-arc,  as  the  unit  of  the  nervous  svstem,  is  itself  a 
combination  of  three  indispensable  })arts  or  elements:  tlie 
receptor,  the  conductor  and  the  effector.  Ty})ically  these 
structures  contain  at  least  four  cells,  one  for  the  receptor, 


164  The  Unity  of  the  Organism 

two  for  the  conductor,  and  one  for"  the  effector.  An  illus- 
tration would  be  a  reflex-arc  consisting  of  a  tactile  cell  of 
a  touch  organ,  the  two  cells  constituting  the  conducting 
path,  and  an  effector  cell  in  a  muscle.  A  point  of  special 
interest  in  connection  with  such  an  arc  is  that  the  dependence 
of  the  parts  on  one  another  is  such,  in  the  highest  develop- 
ment of  the  arc,  that  the  specific  action  of  each  part  is 
dependent  on  the  specific  and  connected  action  of  the  other 
parts.  "The  optic  nerve  itself,"  to  quote  Sherrington,  "is 
unable  to  enter  into  a  heightened  phase  of  its  own  specific 
activity  on  the  application  of  light.  Initiation  of  nervous 
activity  by  light  is  the  exclusive  (in  this  instance)  function 
of  cells  in  the  retina,  retinal  receptors."  ^  And  of  course 
without  brain  cells  as  effectors  for  vision  the  specific  activi- 
ties of  receptors  and  conductors  would  be  impossible.  Not 
only  would  an  optic  nerve  fiber's  conducting  ability  be  useless 
without  a  retinal  cell  on  the  one  hand  and  a  brain  cell  on  the 
other,  but  the  very  conductive  act  itself  would  not  be  fully 
performed. 

This  specific  dependence,  as  it  might  be  called,  of  the 
parts  of  a  reflex-arc  is  so  significant  that  another  example 
may  be  profitably  noticed.  A  striking  one  is  afforded  by 
the  effects  of  the  passage  of  gall  stones  through  the  gall 
ducts,  instanced  by  Sherrington,  partly  on  the  basis  of  his 
own  studies.  The  excruciating  pains  associated  with  this 
malady  are  due  to  the  distention  of  the  wall  of  the  gall  duct 
by  the  passage  through  it  of  the  mineralized  organic  con- 
cretions which  constitute  the  gall  stones.  Tlie  point  in 
this  for  us  is  the  fact  that  though  the  stimulus  which  pro- 
duces the  pain  is  mechanical  and  acts  upon  the  wall  of  the 
duct,  this  stimulus  is  so  peculiar  that  other  sorts  of  me- 
chanical stimuli  of  the  same  tissue,  even  to  cutting  and 
wounding,  give  no  sense  of  pain.  Though  the  duct  may 
be  cut  without  causing  pain,  pain  may  be  produced  by 
injecting  the  duct  to  distention  with  neutral  fluid.    "Marked 


Neural  Integration  1()5 

reflex  effects  can  be  excited  [by  fluid  1  from  the  very  or£rans 
the  cutting  and  wounding  of  which  remains  witliout  ett'ect."  ^ 
The  reflex  mechanism  inyolyed  is  adjusted  to  respond  only 
to  a  stimulus  of  a  special  kind.  In  tliis  it  is  comparable  to 
the  optical  mechanism   referred  to  aboye. 

These  illustrations  show  something  of  the  general  nature 
of  the  reflex-arc  as  an  integrated  structure.  But  we  want 
to  know  something  about  the  part  contributed  by  the 
difl*erent  constituents  of  the  arc  to  this  nature.  The  spe- 
cific office  of  sense  organs  yiewed  as  the  receptor  members  of 
the  arcs  will  first  engage  attention.  "The  main  function  of 
the  receptor  is  therefore  to  lower  the  threshold  of  excit- 
ahility  of  the  arc  for  one  kind  of  stimulus,  and  to  heighten 
it  for  all  others.'^  ^  This  principle  is  so  important  that  we 
must  allow  no  yagueness  as  to  its  meaning.  It  means  that 
while  more  than  one  sort  of  stimulus  may  put  a  particular 
sense  apparatus  into  operation,  there  is  one  and  only  one 
kind,  namely,  that  to  which  the  sense  organ  is  naturally 
subject,  that  elicits  the  reflex  in  its  normal  or  type  expres- 
sion. A  sense  organ  may  be  looked  upon  as  a  group  of 
receptors  attuned  to  a  special  stimulus,  as  contrasted  with 
that  of  the  general  stimuli  to  which  an  organism  is  always 
subject  by  being  always  in  contact  with  its  enyironment. 

In  illustration  Sherrington  instances  the  fact  that  the 
threshold  for  the  touch-sensation  is  lower  for  a  mechanical 
stimulus  than  for  an  electrical  stimulus.  Haying  regard  to 
the  whole  lot  of  reflex-arcs  of  the  body,  we  may  say  that 
the  different  sense  organs  constitute  mechanisms  of  selective 
excitability/  for  the  different  stimuli,  each  organ  })elng  so 
adapted  to  its  natural  stimulus  that  it  responds  to  this 
better  than  to  any  other.  This  reference  to  the  sense  or- 
gans as  adaptive  makes  it  desirable  to  notice  the  fact  that, 
according  to  Sherrington,  electricity  is  neyer  an  ade(ju<ite 
stimulus  of  any  sense  organ  because  it  is  not  a  natural 
stimulus   for  any,   since  it   does   not  enter  into   the   natural 


166  The  Unity  of  the  Organism 

environment  of  the  organism.  This  is  important  in  view 
of  the  large  use  made  of  electricity  in  experimental  work 
on  nerve  physiology.  To  what  extent  may  accepted  gen- 
eralizations concerning  various  reactions  be  influenced  by 
the  inadequacy  of  the  stimuli  applied.^ 

We  glance  next  at  the  physiological  character  of  the  con- 
ducting member  of  the  arc.  It  is  hardly  necessary  to  say 
that  typically  this  part  is  found  in  the  nerve  cords  every- 
where present  in  the  body.  The  most  general  truth  of  sig- 
nificance for  us  is  the  fact  that  having  regard  to  conduction 
through  the  arc  as  a  whole,  the  process  is  different  in  many 
important  respects  from  conduction  in  the  nerve  cord  alone. 
Conduction  in  the  arc  as  an  integrated  whole  is  quite  dif- 
ferent from  conduction  in  the  part  of  the  arc  specially 
devoted  to  that  office.  To  illustrate  by  a  quotation:  "An- 
other remarkable  difference  between  reflex-arc  conduction 
and  nerve-trunk  conduction  is  the  irreversibility  of  direc- 
tion in  the  former  and  the  reversibility  of  the  latter."  ^ 
This  is  only  another  way  of  expressing  the  well-established 
fact  that  nerve  impulses  are  incapable,  in  the  higher  organ- 
isms especially,  of  running  in  more  than  one  direction. 
Sensory  excitations  can  go  only  centripetally  and  motor  ex- 
citations only  peripherally.  As  Sherrington  points  out,  this 
is  part  of  the  "law  of  forward  direction"  of  the  neural  system. 
It  is  fully  established  that  nel'^'e  impulses  may  run  in  both 
directions  when  a  stimulus  is  applied  at  any  point  or  a 
nem-e  trunk,  whether  the  trunk  be  motor  or  sensory.  Taken 
by  itself  this  fact  might  encourage  a  somewhat  careless  ob- 
server to  think  of  the  nervous  s^'Stem  as  more  or  less  hit-or- 
miss  in  its  structure  and  action,  it  being  able  to  work  one 
way  as  well  as  another,  the  final  result  being  determined  by 
where  the  stimuli  happen  to  be  applied.  As  a  matter  of 
fact,  though,  when  we  come  to  consider  the  real  unit  of 
neural  organization,  the  reflex-arc,  instead  of  any  of  the 
constituents  of  that  arc  taken  separately,  all  suggestion  of 


Neural  Iniegration  IHT 

hapliazardness  disappears.  Nerve  conduction  in  a  nerve 
trunk  as  an  isolated  phenomenon  has  neither  existence  nor 
meaning  for  the  actual  organism.  ^J'he  conchicting  element 
par  excellence  of  the  real  unit  of  nerve  organization  is 
differentiated  with  reference  to  the  otlier  elements  of  that 
unit,  and  normally  acts  only  in  such  relation.  The  circum- 
stance that  this  element  is  found  to  liave  tlie  ahihty  to  act 
somewhat  differently  under  different  but  abnormal  rela- 
tions signifies  little  or  nothing,  so  far  as  one  can  see,  for 
the  normal  workings  of  the  nervous  system.  What  it  does 
show  is  something  of  the  diversity  and  plasticity  of  tlie 
organism  because  of  its  latent  abilit}-  to  act  otherwise  tlian 
in  nature  it  does  act  when  conditions  are  imposed  upon  it 
which  are  wholly  new  to  it. 

Although  the  explanation  of  this  irreversibility  of  direc- 
tion appears  not  to  be  known  vnih.  certainty,  the  suggestion 
that  it  may  be  connected  with  a  difference  in  penneabUity 
of  the  synaptic  membrane  between  cells  of  the  reflex-arc  to 
certain  ions,  depending  upon  which  side  of  tlie  membrane  is 
presented  to  these  ions,  is  plausible  and  indicates  the  re- 
sourcefulness, as  one  might  say,  of  structural  and  functional 
method  by  which  the  organization  of  the  living  being  is  ac- 
complished. Should  this  suggestion  prove  to  be  correct, 
the  question  would  arise,  how  comes  it  that  the  synaptic 
membrane  is  thus  differential  in  its  action.'^  And  no  ade- 
quate answer  would  be  forthcoming  that  did  not  take  cogni- 
zance of  the  fact  that  the  membrane  assumed  this  differential 
mode  of  acting  as  part  and  parcel  of  the  differentiation  and 
integration  as  the  reflex-arc  as  a  whole. 

Sherrin£rton  deals  with  a  whole  series  of  other  differences 
between  nerve  trunk  conduction  and  reflex-arc  conduction, 
such  as  the  phenomena  of  summation  of  stinnili,  that  is, 
the  adding  up  of  excitations  too  slight  taken  singly  to  pro- 
duce reaction,  until  the  aggregate  brings  response.  Such 
are  rhythmic  activity  in  response  to  stimuli  which  are  not 


168  The  Unity  of  the  Organism 

rhythmic,  or  at  least  not  of  the  same  time-intervals  as  the 
excitations,  gradation  in  intensity  of  reaction,  and  so  on. 
To  these  he  devotes  more  than  a  hundred  pages,  all  under 
the  common  heading,  "Coordination  in  the  Simple  Reflex." 
It  is  unnecessary  to  follow  the  matter  further,  sufficient 
having  been  given  to  show  something  of  the  variety  of  ends, 
all  definitely  and  specifically  contributory  to  the  needs  of 
the  organism  as  such,  and  all  accomplished  through  differ- 
entiation and  integration  of  one  relatively  simple  organic 
structure,  the  reflex-arc. 

The  Integration  of  Reflex-Arcs 

Our  next  task  is  to  learn  something  of  the  combinations 
among  the  myriads  of  reflex-arcs  of  which  the  higher  nerv- 
ous system  is  composed.  The  definition  of  the  reflex-arc 
given  on  a  preceding  page  presents  this  mechanism  in  its 
simplest  terms,  and  so  far  as  the  definition  indicates  it  would 
seem  possible  for  it  actually  to  exist  and  be  operative  in 
this  simplicity.  In  fact,  physiologists  are  accustomed  to 
assume  such  an  entity  and  to  call  it  the  simple  reflex-arc, 
and  its  activity  the  simple  reflex.  But  while  such  a  concep- 
tion is  convenient  and  helpful  for  didactic  purposes,  espe- 
cially as  an  aid  to  visualizing  the  earliest  stage  of  cell  inte- 
gration in  the  evolution  of  the  nervous  system,  as  a  matter 
of  fact,  according  to  Sherrington,  it  probably  never  exists 
in  nature.  "A  simple  reflex  is  probably  a  pure  abstract 
conception,  because  all  parts  of  the  nervous  system  are 
connected  together  and  no  part  of  it  is  probably  ever 
capable  of  reaction  without  affecting  and  being  affected  by 
various  other  parts,  and  it  is  a  system  certainly  never  abso- 
lutely at  rest."  ^ 

The  factual  basis  for  the  conception  that  one  of  the  most 
essential  distinctions  between  the  nervous  system  and  other 
systems  of  the  body  is  the  integratedness  of  the  former,  is 


Neural  Integration  1()9 

well  expressed  bj  Donaldson :  "Isolated  groups  of  nerve- 
cells  do  not  occur.  Indeed,  a  group  of  nerve-cells  discon- 
nected from  the  other  nerve-tissues  of  the  body,  as  nuiscles 
or  glands  are  disconnected,  would  be  witliout  j)hvsiological 
significance.  It  is  desirable,  therefore,  to  emphasize  the 
fact  that  by  dissection  the  nervous  system  is  found  to  l)o 
connected  throughout  its  entire  extent."  '' 

I  would  ask  the  reader  to  consider  these  statements  in  the 
light  of  the  cellular  conception  of  the  organism  maintained 
in  this  volume  and  especially  dwelt  upon  in  the  chapters  on 
the  cell-theory,  namely,  that  the  true  way  of  viewing  the 
organism  is  not  as  being  built  up  of  cells  in  the  sense  of 
having  been  constructed  by  the  bringing  together  of  pre- 
viously isolated  cells,  as  a  brick  house  is  built  up  of  bricks ; 
but  rather  as  being  composed  of  cells  through  resolving 
itself  into  these  as  it  increases  in  size  and  differentiates 
itself  into  its  organs  and  functions.  With  such  a  concep- 
tion Sherrington's  formulation,  based  on  the  findings  of 
vast  observational  and  experimental  research  on  the  struc- 
ture and  action  of  the  nervous  system,  is  in  perfect  accord. 
If  we  see  in  the  completed  nervous  system  a  complex 
mechanism  developing  as  a  unit  subservient  at  all  stages  to 
the  needs  of  the  organism  as  a  whole,  the  myriads  of  reflex- 
arcs  then  present  themselves  as  final,  as  end-stages  in  the 
differentiation  and  not  as  initial  states ;  and  the  universal 
organic  and  functional  connection  with  one  another,  affirmed 
in  the  quotation,  would  be  just  what  we  might  expect. 

If,  on  the  other  hand,  the  integral  nervous  system 
were  built  up  in  a  literal  sense,  that  is,  by  the  actual  coming 
into  connection  with  one  another  of  previously  isolated 
simjole  reflex-arcs,  such  arcs  ought  to  be  demonstrable  botli 
in  the  ontogeny  of  higher  animals  and  in  the  adults  of  the 
lower  metazoa. 

No  one  should  be  beguiled  into  the  notion  that  the  readily 
observed  facts  of  ontogeny  of  the  nervous  system,  the  vari- 


170  The  Unity  of  the  Organism 

ous  processes,  dendrites  and  axones,  do  actually  grow  out 
on  nerve  cells  and  bring  cells  into  connection  with  one 
another  and  with  receptor  and  effector  cells,  and  that  a 
functional  coordination  is  thus  finally  reached  does  not 
exist  in  any  way  or  degree  in  the  early  stages.  Proof  that 
the  completed  stages  of  neural  integration  cannot  be  ac- 
complished without  the  production  of  cell-growths  which 
put  cells  into  connection  with  one  another  is  a  very  different 
thing  from  proof  that  these  cells  were  once  wholly  isolated 
from  one  another,  and  that  the  outgrowths  which  establish 
the  final  connections  were  initiated  by  impulses  which  origi- 
nated wholly  within  the  cells ;  as,  by  way  of  illustration,  the 
branches  of  two  young  trees  standing  not  far  apart  might 
come  into  contact  as  the  trees  increased  in  size.  It  is  a 
matter  of  elementary  knowledge  of  animal  development  that 
in  this  sense  the  cells  of  the  nervous  system  are  never  iso- 
lated either  from  other  nerve  cells  or  cells  of  certain  other 
parts  of  the  organism.  Until  we  know  vastly  more  than 
we  do  know  of  the  chemical  nature  of  intercellular  substances 
and  of  the  chemical  and  physical  activities  which  go  on  at 
the  planes  of  contact  between  cells,  nothing  could  be  more 
gratuitous  and  unscientific  than  to  assume  that  nerve  cells 
differentiate  as  they  would  were  they  not  in  some  measure 
vitally  associated  with  one  another  from  the  very  begin- 
ning. Indeed,  such  positive  knowledge  as  we  have  tends 
strongly  against   such  an  assumption. 

Recall,  for  example,  the  fundamental  part  chemical  mes- 
sengers play  in  development.  And  Sherrington's  insistence 
on  the  role  of  intercellular  substance  and  "surfaces  of  sepa- 
ration" between  cells  in  the  functioning  of  the  adult  nervous 
system  is  much  to  the  point  for  this  contention.  Intercellu- 
lar as  well  as  intracellular  conduction  must  be,  he  main- 
tains, expected  in  the  reflex-arc  on  the  basis  of  the  cell 
theory.^ 


Neural  Integration  171 

The  Spreading  and  Compoiindijig  of  Reflexes 

As  a  practical  matter  study  of  tlie  intcfrratlvL'  action  of 
the  nervous  systems  never  gets  away  from  integration  among 
reflex-arcs    and   reflexes    any   more   than    it   docs    from    tlie 
cellular   intergration    fundamental    to    the    reflex-arc    itself. 
The  general  nature  of  the  study  always  involves  the  "spread 
of  reflexes  over  a  wide  range  of  nervous  arcs."     The  idea 
is  more  fully  stated  in  the  following:  "This  compounding  of 
reflexes   with   orderliness   of   coadjustment  and   of  sequence 
constitutes    coordination,    and   want    of   it    inco-ordination. 
We   may    therefore    in    regard    to    coordination   distinguish 
coordination  of  reflexes  simultaneously  proceeding,  and  co- 
ordination  of   reflexes   successively   proceeding.      The   main 
secret   of  nervous   coordination  lies   evidently   in   the   com- 
pounding of  reflexes."  ^    For  the  rest,  all  we  can  do  or  need 
do  to  meet  the  requirements  of  this  discussion  is  to  get  at 
the  main  principles  as  illustrated  by  examples  of  this  com- 
pounding of  reflexes. 

As  a  starting  point  for  the  discussion  of  this  larger 
aspect  of  neural  integration,  Sherrington  takes  what  he  has 
called  "the  principle  of  the  common  path."  Basal  to  this 
conception  is  the  famihar  fact  that  a  stimulus  applied  at  a 
single  point,  on  the  surface  of  the  body,  for  example,  gives 
rise  to  a  nerve  impulse  which  may  reach  a  great  number 
of  muscles  or  glands.  A  single  receptor  with  its  neurone 
must  be  in  communication  with  a  great  number  of  effectors, 
some  of  which  are  very  diff'erent  in  kind.  Looking  at  the 
arcs  from  the  efl'ector  ends,  as  one  may  say,  it  is  obvious 
that  the  impulses  reaching  innumerable  eff'ectors  must  come 
over  a  single  conducting  course. 

On  the  other  hand,  since  a  given  muscle  or  gland  may  be 
reached  by  impulses  from  a  great  number  of  reflex-arcs, 
the  efl*ector  and  its  neurone  must  })e  the  "common  patli" 
for  all  these  impulses,   often   very   different   in   quality   and 


172  The  Unity  of  the  Organism 

source.  "While  the  receptive  neurone  forms  a  private  path 
exclusively  serving  impulses  of  one  source  only,  the  final 
or  efferent  neurone  is,  so  to  say,  a  public  path,  common 
to  impulses  arising  at  any  of  many  sources  of  reception."  ^ 

An  example  which  illustrates  the  general  principle  of  the 
common  path,  and  several  phenomena  incidental  to  this, 
and  one  which  has  been  much  investigated  by  Sherrington, 
is  that  of  the  scratching  reflex  in  dogs.  It  has  long  been 
known  that  in  several  land  vertebrates  which  have  the 
habit  of  scratching  the  side  and  back  of  the  forward  part 
of  the  body  with  the  hind  foot,  the  scratching  movement 
may  be  elicited  by  appropriate  stimuli  applied  to  the  area 
reached  by  the  foot  as  a  pure  reflex ;  that  is,  in  the  absence 
of  any  chance  for  impulses  from  the  brain  to  reach  the 
parts  involved  in  the  activities.  Since  the  scratching  move- 
ment consists  in  bringing  the  hind  leg  forward  and  upward, 
and  for  scratching,  a  rhythmic  movement  of  the  foot,  the 
muscles,  both  flexor  and  extensor,  of  the  thigh,  leg  and 
foot  must  be  involved.  And  since  the  reflex  can  be  induced 
by  a  stimulus  applied  at  any  point  within  the  large  recep- 
tive field  (i.  e.,  nearly  the  whole  side  and  back  of  the  body), 
impulses  started  from  various  parts  of  the  field  must  pass 
through  one  and  the  same  neurone  in  the  muscles  concerned. 

And  here  comes  in  a  fact  showing  another  aspect  of  the 
integration  of  reflexes  in  this  case.  A  stimulus  at  a  given 
point  in  the  field  too  weak  by  itself  to  elicit  the  reflex  may 
bring  it  on  when  acting  in  combination  with  weak  stimuli 
at  other  points  in  the  field.  Sherrington  calls  reflexes 
which  act  together  in  this  way  allied  reflexes. 

Still  another  kind  of  combination  of  reflexes  involving 
the  common  path  principle,  even  more  significant  than  allied 
reflexes,  are  what  are  known  as  proprio-ceptive  reflexes. 
The  kernel  of  this  class  of  reactions  is  the  existence  of  re- 
ceptors in  the  deep  tissues  of  the  body,  that  is,  not  belonging 
to  the  surface  and  hence  not  subject  to   stimuli  from  the 


Neural  Integration  173 

external  world,  but  because  of  their  situation  subject  onlv 
to  stimuli  "given  by  the  orrranisni  itself."  Spcciticallv  the 
sources  of  the  stimuli  are  the  muscles,  joints,  blood  vessels 
and  so  on,  which  by  the  regular  activities  are  always  in 
more  or  less  movable  contact  with  one  another  and  with 
other  parts  and  organs.  Stimuli  from  the  environment 
acting  upon  the  skin  receptors  give  rise  to  reflexes  which 
put  muscles  into  activity,  and  these  activities  serve  in  turn 
as  stimuli  to  receptors  in  the  deep  parts;  and  the  impulses 
arising  from  these  deep  receptors  may  pass  to  still  other 
muscles  over  the  same  effector  neurons  used  bv  im})ulses 
coming  from  skin  receptors.  The  proprio-ceptive  field  is 
a  sort  of  relay  and  supplement  and  extension  of  the  field 
of  contact  with  the  environment,  or  extero-ceptive  field. 

A  simile  used  by  Sherrington  helps  us  to  understand  the 
import  of  this  compounding  of  reflexes :  "The  receptor 
system  bears,  therefore,  to  the  efferent  paths  the  relation 
of  the  wide  ingress  of  a  funnel  to  the  narrow  egress.  Fur- 
ther, each  receptor  stands  in  connection  not  with  one  efferent 
only  but  with  many — perhaps  with  all — though  as  to  some 
of  these  only  through  synapses  of  high  resistance.  The 
simile  to  a  funnel  will  therefore  be  bettered  by  supposing 
that  within  the  general  systemic  funnel,  of  which  the  base 
is  five  times  wider  than  the  egress,  the  conducting  ])aths 
from  each  receptor  may  be  represented  as  a  funnel  Inverted 
so  that  its  wider  end  is  more  or  less  coextensive  with  the 
whole  plane  of  emergence  of  the  final  connnon  patiis.  This 
gives  some  idea  of  the  enormous  formation  of  common 
paths  from  tributary  paths  which  must   take  ])lace.'''  ^"^ 

And  Sherrington  forces  home  the  truth  of  the  scope  o/ 
the  combinations  by  calling  attention  to  the  fact  that  under 
strychnine  poisoning  "a  muscle  can  be  excited  from  prac- 
tically any  afferent  nerve  in  the  body."  This  is  equiva- 
lent to  saying,  he  remarks,  "that  each  final  connnon  ])ath  is 
in  connection  with  jjractically  each  one  of  all  the  receptors 


174  The  Unity  of  the  Organism 

of  the  body."  11 

Even  though  this  statement  should  prove  to  be  too  strong, 
it  certainly  contains  truth  enough  to  show  "the  profusion 
in  which  common  paths  exist." 

The  functional  side  of  the  total  reflex  system,  the  struc- 
tural side  of  which  was  visualized  by  the  funnel  simile  stated 
above,  may  be  regarded  as  pretty  well  presented  by  certain 
phenomena  known  as  irradiation.  What  is  meant  by  this 
is  quite  clear,  in  its  general  outlines,  from  the  following: 

"The  more  intense  the  spinal  reflex  .  .  .  the  wider,  as  a 
general  rule,  the  extent  to  which  the  motor  discharge  spreads 
around  its  focal  area.  Thus,  as  stimulation  of  the  planta 
causing  the  flexion-reflex  is  increased  there  is  added  to  the 
flexion  of  the  homonymous  hind  limb  extension  of  the  crossed 
hind  limb,  then  in  the  homonymous  fore  limb  extension  at 
elbow  and  retraction  at  shoulder,  then  at  the  crossed  fore 
limb  flexion  at  elbow,  extension  at  the  wrist,  and  some 
protraction  at  shoulder ;  also  turning  of  the  head  toward 
the  homonymous  side,  and  often  opening  of  the  mouth, 
also  lateral  deviation  of  the  tail.  According  to  circum- 
stance, especially  according  to  intensity  of  stimulation,  the 
field  of  end-eff*ect  of  the  flexion-reflex  may  vary  from  a 
minute  field  occupying  part  of  a  flexor  muscle  of  the  knee 
to  a  field  including  musculature  in  all  four  limbs  and  neck 
and  head  and  tail."i^ 

Antagonistic   Reflexes   in   Skeletal  Muscle   Groups   Finally 

Integrative 

So  far  what  has  been  said  about  the  interaction  between 
reflexes  has  dealt  only  with  interactions  which  are  har- 
monious with  one  another  in  various  modes  of  combination. 
But  there  are  antagonistic  or  competitive  as  well  as  har- 
monious interactions.  These  must  now  receive  attention. 
As  an  illustration  we  make  use  again  of  the  scratching  re- 


Neural  Integration  175 

flex.  "If,"  says  Sherrington,  "while  stiniuhitloii  of  tlie  skin 
of  the  shoulder  is  evokinfr  the  scratch-reflex,  the  skin  of 
the  hind  foot  of  the  sanw  side  is  stimulated,  the  scratching 
may  be  arrested."  ^^ 

Then  the  author  proceeds  to  show,  by  description  and 
diagrams,  how  the  two  excitations  here  involved,  one  from 
the  skin  of  the  side  of  the  body,  the  other  from  the  hind 
foot  of  the  same  side,  have  the  "same  final  common  i)ath" 
to  the  muscles  of  the  hip  and  leg,  which,  however,  they  "w,?^ 
to  different  effect'^',  that  is,  the  one  to  excite,  the  other  to 
inhibit,  contraction  of  the  muscles  concerned. 

At  once  there  arises  the  important  question :  What  is  the 
meaning  of  such  antagonisms,  such  seeming  want  of  har- 
mony, as  tliis.'^  How  deep-seated  is  the  competition  thus 
shown  .'^  Does  it  amount  to  an  "ultimate  truth"  in  the  na- 
ture of  the  organism,  thereby  furnishing  an  argument  in 
favor  of  the  "struggle  of  the  parts"  as  an  explanation  of 
the  organism?  Or  is  it  possible  that  the  antagonism  is 
secondary  to  the  wider  needs  and  activities  of  the  organism 
as  a  whole.? 

A  partial  answer  to  this  inquiry  is  found  in  the  character 
of  the  stimuli  operative  in  the  two  opposing  fields.  "Stimu- 
lation of  the  skin  of  the  hind  foot  by  any  of  the  various 
stimuli  that  have  the  character  of  threatening  the  part  with 
damage  causes  the  leg  to  be  flexed,  drawing  the  foot  up 
by  steady  maintained  contraction  of  the  flexors  of  the  ankle, 
knee  and  hip."  ^"^ 

In  other  connections  Sherrington  dwi-lls  on  the  peculiari- 
ties of  efl'ect  and  importance  from  the  standpoint  of  adap- 
tation, of  stimuli  of  this  class,  as  for  example  ])ricking, 
strong  squeezing,  injurious  heat,  and  so  on. 

From  the  descriptions  we  notice  that  the  actions  of  the 
hind  leg  involved  in  the  scratching-rcflex  arc  considrrably 
diff*erent  from  those  involved  in  withdrawing  the  hind  foot 
from  stimuli  of  liarmful  portent,      l^ut  since  the  same  nnis- 


176  The  Unity  of  the  Organism 

cles  are  necessarily  involved  in  the  two  sorts  of  action,  and 
since  the  stimuli  reaching  these  muscles  from  the  two 
sources  must  use  the  same  final  paths  to  those  muscles,  back 
of  the  antagonism  between  the  two  sets  of  reflexes  is  the 
question,  which,  in  a  given  case,  is  more  important  to  the 
organism,  the  scratching  action  or  the  withdrawal-from- 
danger  action? 

Since  the  scratch-reflex  in  the  dog  is  probably  connected 
primarily  with  flea  and  other  insect  bites,  and  since  on  the 
whole  it  may  be  assumed  that  these  are  rather  annoyances 
than  real  dangers  to  life  and  limb  as  the  "nocuous"  excita- 
tions are  by  fundamental  nature,  it  would  be  fair  to  infer 
that  though  either  reflex  might  under  certain  circumstances 
inhibit  the  other,  the  threshold  for  the  injury-escaping  re- 
flex would  be  lower.  I  do  not  know  that  there  is  anything 
in  the  evidence  which  bears  directly  on  this  point,  but  the 
question  is  one  that  would  surely  arise  were  the  whole  sub- 
ject of  antagonistic  reflexes  to  be  looked  at  from  the  stand- 
point of  the  needs  and  adaptations  of  the  normally  living 
organism. 

The  way  certain  other  reflexes,  antagonistic  in  a  sense, 
are  yet  correlated  in  a  larger  sense,  is  more  obvious  than  in 
the  case  just  given.  Thus  the  reflex  appertaining  to  two 
limb  muscles  which  oppose  each  other  does  not  merely  ac- 
tivate the  muscle  which  contracts ;  it  simultaneously  causes 
depression  of  the  opposing  muscle.  Cooperative  antagon- 
ism, as  it  might  be  called,  of  this  general  sort  is  widespread 
among  higher  animals,  and  applies  to  glandular,  circulatory 
and  various  other  mechanisms  as  well  as  to  the  muscular. 
Indeed,  it  being  undoubtedly  true,  as  Sherrington  repeatedly 
points  out,  that  the  "outward  behavior"  of  animals  involves 
a  great  variety  of  movements  which  proceed  in  an  orderly 
sequence,  if  they  are  normal,  it  seems  almost  necessary  to 
suppose  that  really  all  normal  reflexes  must  be  cooperative 
and  harmonious  with  reference  to  the  organism  as  a  whole, 


Neural  Integration  177 

even  though  when  viewed  one  bj  one  or  in  secondary  groups 
they  are  antagonistic. 

Such  a  conception  of  the  real  nature  of  antagonistic  re- 
flexes is  favored  by  the  seemingly  general  fact  tliat  these 
reflexes  are  seldom  if  ever  really  destructive  of  one  aiiotlur, 
since  they  do  not  act  upon  one  another  simidtaneouslif. 
Their  antagonism  consists  in  a  successional  opposition  to 
one  another.  As  they  follow  one  another  one  acts  in  the 
opposite  direction  to  the  other,  and  the  antagonism  is  tlie 
more  real  in  that  frequently  tliey  overlaj)  to  some  extent. 
But  as  already  said  this  overlapping  probably  never  amounts 
to  complete  coincidence.  Such  overlappings  and  other 
forms  of  partial  opposition  constitute  the  phenomena  of  in- 
hibition which  play  a  great  role  in  the  sum  total  of  reflexes 
of  the  organism.  This  is  part  of  tlie  method  by  whlcli 
transition  is  accomplislicd  from  one  reflex  to  anotlier,  where 
the  same  muscles,  for  example,  execute  both,  l^iit  tlie  fac-t 
that  the  transition  is  accomplished  normally  "without  con- 
fusion," to  use  Sherrington's  phrase,  shows  the  subordina- 
tion of  the  inhibitions  to  the  organism  as  a  whole. 

Another  important  fact  to  which  Sherrington  calls  at- 
tention is  that  inhibitions  which  reflexes  produce  upon  one 
another  never,  so  far  as  is  known,  result  in  injury  to  the 
tissues  involved.  Genuine  opposition  of  reflexes,  as  of  any 
other  sorts  of  physical  or  chemical  action,  would,  according 
to  all  our  conceptions  of  natural  bodies,  have  deleterious 
eff'ects  on  the  opposing  bodies.  As  a  matter  of  fact,  in- 
hibiting reflexes  not  only  do  not  injure  the  mechanisms  in- 
volved, but  actually  prepare  them  for  greater  fimetional 
activity  later  on.^^ 

This  beneficent  efl'ect,  as  it  might  be  called,  of  inhibition 
is  perhaps  illustrated  by  certain  forms  of  compensatory 
reflexes.  Thus,  stinuilation  of  the  central  end  of  the  nerve 
to  the  extensor  muscles  of  the  dog's  knee  results  in  contrac- 
tion of  the  flexors  of  the  hip  and  knee.     But  on  removal  of 


178  The  Unity  of  the  Organism 

this  stimulus  contraction  of  the  extensor  muscles  imme- 
diately succeeds,  this  "rebound"  being  especially  marked 
in  the  vasto-criireus  muscle,  the  main  knee  extensor. -^^ 

The   Antagonisms    within    the    Autonomic    System   Finally 

Integrative 

It  was  mentioned  above  that  "cooperative  antagonism"  in 
nervous  action  is  widespread  in  the  animal  mechanism.  The 
illustrations  given  pertained  to  the  cerebrospinal  system 
and  the  innervation  of  skeletal  muscles.  We  must  now  fol- 
low this  subject  farther  and  deal  with  the  somewhat  similar 
phenomena  presented  by  the  autonomic  nervous  system. 
This  subject  was  treated  to  some  extent  in  the  chapter  on 
internal  secretions,  to  the  extent,  that  is,  that  it  implicated 
the  endocrinal  glandular  system.  The  presentation  here 
will  involve  some  repetition  of  what  was  said  in  the  previous 
discussion,  but  the  nature  and  importance  of  the  subject 
will  justify  this. 

Cannon  appears  to  have  been  the  first  to  make  clear  the 
similarity  between  the  opposing  action  of  the  subdivisions  of 
the  autonomic  nervous  system  and  what  Sherrington  calls 
the  reciprocal  innervation  of  antagonistic  skeletal  muscles. 
"As  the  above  description  has  shown,"  Cannon  writes,  "there 
are  peripheral  oppositions  in  the  viscera  corresponding  to 
the  oppositions  between  flexor  and  extensor  muscles."  ^^ 

The  description  referred  to  is  summed  up  in  the  state- 
ment that  many  of  the  viscera  and  other  parts  of  the  body 
are  innervated  by  either  the  cranial  or  sacral,  i.e.,  the  ter- 
minal autonomics,  and  also  by  the  thoracico-lumbar  or  mid- 
dle autonomic,  this  double  innervation  being  such  as  to  be 
statable  thus:  ''When  the  mid-part  meets  either  end-part  in 
any  viscus  their  effects  are  antagonistic.^'  '^'^  The  heart- 
beat is  slowed  by  the  cranial  autonomic  and  quickened  by  the 
thoracico-lumbar.       Contraction    of    the    smooth    muscular 


Neural  Integration  179 

layer  of  the  stomach  and  small  intestine  to  produce  '*tone"  is 
increased  on  the  whole  by  the  cranial  autonomic,  while  gas- 
tric peristalsis  and  secretion  are  inliibitcd  and  tlie  arterioles 
of  these  organs  are  contracted  by  the  thoracico-lumbar.''* 
The  pupil  of  the  eye  is  contracted  by  the  cranial  autonomic 
and  dilated  by  tlie  thoracico-lumbar.  The  lower  part  of  the 
large  intestine  is  contracted  by  the  sacral  autonomic  and  is 
relaxed  by  the  thoracico-lumbar.  Tlie  discharge  tube  of 
the  urinary  bladder  is  relaxed  by  the  sacral  and  contract j-d 
by  the  thoracico-hmibar.  Tlie  blood  vessels  of  the  erectile 
tissue  of  the  external  genitals  are  dilated  by  the  sacral 
autonomic  and  contracted  by  the  thoracico-lumbar;  and 
so  on. 

Now  it  is  especially  important,  as  Cannon  says,  to  notice 
the  kind  of  service  these  subdivisions  of  the  autonomic  sys- 
tem perform  for  the  organism.  On  considering  the  func- 
tions of  the* cranial  division,  one  recognizes  that  they  have 
to  do  with  bodily  conservation.  "By  narrowing  the  pupil 
of  the  eye  they  shield  the  retina  from  excessive  light.  By 
slowing  the  heart  rate,  they  give  the  cardiac  muscle  longer 
periods  for  rest  and  invigoration.  And  by  j)roviding  for  the 
flow  of  gastric  juice  and  by  supplying  the  muscular  tone 
necessary  for  contraction  of  the  alimentary  canal,  thoy 
prove  fundamentally  essential  to  the  processes  of  proj)er 
digestion  and  absorption  by  which  energy-^delding  material 
is  taken  into  the  body  and  stored.  To  the  cranial  division 
of  the  visceral  nerves,  therefore,  belongs  the  quiet  service 
of  building  up  reserves  and  fortifying  the  body  against  times 
of  need  or  stress."  ^^ 

Pasing  to  the  sacral  division,  one  sees  that  as  concerns 
its  distribution  to  the  digestive  and  urinary  viscera,  its  of- 
fice is  that  of  accomplishing  the  discharge  of  refuse  material 
from  the  body.  Hence,  *'like  the  cranial  division,  the  sacral 
is  engaged  in  internal  service  to  the  body,  \n  the  ])orfonn- 
ance  of  acts  leading  immediately  to  greater  comforf."-'^ 


180  The  Unity  of  the  Organism 

So  much  in  illustration  of  the  service  of  the  two  end  di- 
visions of  the  autonomic  nervous  system. 

What,  exactly,  we  now  inquire,  is  the  nature  of  the  service 
performed  by  the  thoracico-lumbar  division?  Since  this 
part  is,  as  already  seen,  antagonistic  in  its  action  to  both 
the  cranial  and  sacral  parts  wherever  it  innerv^ates  an  organ 
or  part  also  innerA-ated  by  these  end-parts,  the  inference  is 
readily  drawn  that  its  service  to  the  organism  as  a  whole 
would  also  be  in  a  sense  opposed  to  the  services  of  the 
cranial  and  sacral  divisions.  Only  a  moment's  reflection 
is  necessary  to  recognize  that  at  some  points  at  least  this 
is  so.  For  instance,  quickening  of  the  heart  beat  through 
innervation  by  the  thoracic  autonomic  nerves,  thereby  send- 
ing the  blood  stream  more  rapidly  and  strongly  through  the 
whole  body,  is  clearly  opposed,  so  far  as  these  acts  in  them- 
selves are  concerned,  to  the  slowing  of  the  heart  and  hence 
of  the  blood  stream  by  the  vagus  nei'A^e,  i.e.,  by.  cranial  au- 
tonomic innervation.  Now  this  can  mean  nothing  else  than 
that  whereas  the  vagal  (cranial)  autonomic  action  is  in 
the  interest  of  upbuilding  and  conserving  the  organism  in 
its  whole  normal,  wonted  life,  the  speeded-up  action  through 
the  thoracic  autonomic  is  in  the  interest  of  some  special, 
more  or  less  temporary  need  or  condition.  But  obvious  and 
important  as  is  this  opposition  between  the  two  subdivisions 
as  concerns  heart  innervation,  of  not  less  importance,  though 
less  easily  observable  in  their  full  scope,  are  the  oppositions 
and  reciprocations  brought  about  in  the  peripheral  blood 
vessels  of  the  whole  body  through  the  innerv^ation  of  the 
muscles  of  the  arterial  walls  by  the  thoracico-lumbar  au- 
tonomic in  connection  with  the  heart  action.  These  in- 
nervations, coupled  with  those  of  the  smooth  muscles  of  the 
gastro-intestinal  canal,  of  the  sweat  and  probably  other 
peripheral  glands,  of  the  smooth  muscles  of  the  hairs  in 
mammals,  of  the  adrenal  medulla  for  the  secretion  of 
adrenin,  and  of  the  liver  for  releasing  stored  carbohydrates 


Neural  Integration  181 

into  the  blood,  by  the  thoracico-lunibar  autonomic,  produce 
an  cquiHbrating  mechanism  of  well-ni^li  inconceivable  com- 
plexity and  delicacy,  the  workings  of  which  are,  however, 
beginning  to  be  revealed  to  us,  both  as  to  details  and  as  to 
rationale  for  the  life  of  the  organism  as  a  wliole. 

The  whole  scheme  is  adjusted,  we  may  say,  for  these  two 
necessities  of  the  organism:  (1)  to  serve  the  organism  in  its 
uninterrupted  and  uninterruptible,  and  therefore  normal, 
growth  and  existence;  and  (2)  to  secure  the  perpetuity  of 
the  organism  through  the  recurring  times  of  special  demand 
and  stress  which  are  inevitable  from  the  external  conditions 
of  life  under  which  all  living  beings  exist. 

The  supreme  significance  of  all  this  from  our  standpoint 
is  that  we  get  a  glimpse  of  the  means  by  which  an  extensive 
and  fundamental  number  of  parts  of  the  organism  are  sub- 
ordinated in  their  special  activities  to  the  special  needs  of 
the  whole.  The  following  from  Cannon's  chapter.  Fatigue 
and  Blood  Pressure,  gives  concreteness  and  some  particu- 
larity to  this  statement:  "In  connection  witli  the  foregoing 
considerations  [of  facts  bearing  on  the  value  of  increased 
arterial  pressure  in  pain  and  strong  emotion]  tlie  action  of 
adrenin  on  the  distribution  of  the  blood  in  the  bodv  is  hitjhlv 
interesting.  By  measuring  alternations  in  the  volume  of 
various  viscera  and  limbs,  Oliver  and  Schiifer  proved  that 
the  viscera  of  the  splanchnic  area — e.g.,  the  spleen,  the 
kidneys,  and  the  intestines — suffer  a  considerable  decrease 
of  volume  when  adrenin  is  administered,  wliereas  the  limbs 
into  which  the  blood  is  forced  from  the  splanchnic  region 
actually  increase  in  size.  The  action  of  the  adrenin  indicates 
the  relative  degrees  of  sympathetic  innervation."  -' 

This  last  sentence  is,  of  course,  what  specially  concerns 
us  in  this  discussion.  Continuing,  and  having  in  view  his 
own  researches  into  the  beneficial  effects  on  fatigued  striated 
muscles  of  adrenin  in  the  blood,  Cannon  says:  "At  times  of 
pain  and  excitement  sympatlietic  discharges,  probably  aided 


182  The  Unity  of  the  Organism 

by  the  adrenal  secretion  simultaneously  liberated,  will  drive 
the  blood  out  of  the  vegetative  organs  of  the  interior,  which 
serve  the  routine  needs  of  the  body,  into  the  skeletal  muscles 
which  have  to  meet  by  extra  action  the  urgent  demands  of 
struggle  or  escape.  But  there  are  exceptions  to  the  general 
statement  that  by  adrenin  the  viscera  are  emptied  of  their 
blood.  It  is  well  known  that  adrenin  has  a  vasodilator,  not 
a  vasoconstrictor,  action  on  the  arteries  of  the  heart;  it  is 
well  known  also  that  adrenin  affects  the  vessels  of  the  brain 
and  the  lungs  only  slightly  if  at  all.  From  this  evidence 
we  may  infer  that  sympathetic  impulses,  though  causing 
constriction  of  the  arteries  of  the  abdominal  viscera,  have 
no  effective  influence  on  those  of  the  pulmonary  and  intra- 
cranial areas  and  actually  increase  the  blood  supply  to  the 
heart.  Thus  the  absolutely  and  immediately  essential  or- 
gans— those  the  ancients  called  the  'tripod  of  life' — the 
heart,  the  lungs,  the  brain  (as  well  as  its  instiTiments,  the 
skeletal  muscles) — are  in  times  of  excitement  abundantly 
supphed  with  blood  taken  from  the  organs  of  less  importance 
in  critical  moments."  And  Cannon  concludes  with  the  very 
pertinent  remark:  "This  shifting  of  the  blood  so  that  there 
is  an  assured  adequate  supply  to  structures  essential  for  the 
preservation  of  the  individual  may  reason abh^  be  inter- 
preted as  a  fact  of  prime  biological  significance."  ^-  Indeed 
so !  Even  more  significant,  I  believe,  than  Cannon  has  shown 
us — as  we  shall  see  in  a  later  chapter.  But  the  limitations 
set  at  the  outset  of  tliis  chapter,  namely  that  of  dealing 
only  with  neural  processes  strictly,  especially  as  these  mani- 
fest themselves  in  reflexes,  permits  us  to  go  no  farther  than 
to  call  attention  to  the  fact  that  while  Cannon  finds  "the 
most  significant  feature"  of  the  reactions  he  has  studied  to 
be  "that  they  are  of  the  nature  of  reflexes — they  are  not 
willed  movements,  indeed  they  are  often  distressingly  be- 
yond the  control  of  the  will,"  ^'^  he  yet  has  coupled  them  in 
the   most   positive  way   with   the   emotions,   especially   with 


Neural  Integration  183 

those  of  fear  and  rage,  and  with  pain.  Otht-rwise  stated,  he 
has  coupled  them,  more  definitely  tlian  tliis  lias  been  done 
before,  with  the  conscious  psychic  life  of  the  orrranisni. 

Concluding  Remarks  on  tlie  Significance  of  Neural  Integra- 
tion for  the  Organismal  Standpoint 

Perhaps  enough  illustration  and  general  discussion  have 
been  presented  to  convince  the  reader  not  only  that  "tlie 
nervous  system  functions  as  a  whole"  -*  but  that  this  func- 
tioning is  strictly  subservient  to  the  needs  of  the  organism 
as  a  whole,  whether  the  normal  individual,  living  normally, 
or  the  normal  individual  living  under  special  stress,  be  con- 
sidered. It  is  hoped  the  reader  will  not  have  failed,  despite 
the  brevity  and  inadequacy  of  the  presentation,  to  perceive 
the  fundamental  truth  that  the  organism's  totality  of  activi- 
ties, executed  to  so  large  an  extent  through  the  agency  of  the 
neural  mechanism,  are  in  turn  subordinate  to  the  needs  of 
the  organism  as  related  to  its  natural  eniiromnent.  This  is 
only  another  way  of  expressing  the  truth,  whicli  has  beconn' 
almost  trite  since  the  idea  of  evolution  has  been  an  essen- 
tial part  of  biological  pliilosophy,  that  all,  or  at  least  by 
far  the  major  part,  of  the  activities  of  organisms  are 
adaptive. 

In  connection  with  no  other  organ-system  than  the  nerv- 
ous does  the  truth  come  out  so  patently  that  the  unity  and 
wholeness  of  the  particular  si/sfe?n,  the  unity  and  wholeness 
of  the  organism,  and  the  adaptiveness  of  the  organism  to  its 
environment  are  bound  together  inseparably;  indeed,  may 
almost  be  called  different  aspects  of  one  and  the  same  tnith 
of  living  beings.  The  very  concej^tion  of  adaptation,  at 
least  as  touching  neural  activity,  sccins  (kpciRk-nt  uj)()n  the 
correlatedness,  the  unitedness,  of  the  differentiated  parts. 
The  conception  of  adaptation,  so  far  at  hast  as  reflexes  are 
concerned,   depends   essentially   on    the   mode   of   relation    of 


184  The  Unity  of  the  Organism 

parts.  It  is  non-existent  except  through  a  particular  form 
of  integration. 

Every  specific  act  of  every  part  of  the  nervous  system 
is  primarily  in  the  interest  of  some  other  part  and  function 
of  the  organism  than  itself. 

This  is  particularly  true  for  all  adaptive  acts  of  the 
system. 

Considerations  of  this  sort  seem  not  only  to  justify,  but 
to  render  necessary  some  such  view  of  purpose  in  the  or- 
ganism as  the  following:  "The  infinite  fertility  of  the  or- 
ganism as  a  field  for  adapted  reactions  has  become  more 
apparent.  The  purpose  of  a  reflex  seems  as  legitimate  and 
urgent  an  object  for  natural  inquiry  as  the  purpose  of  the 
colouring  of  an  insect  or  a  blossom.  And  the  importance  to 
physiology  is,  that  the  reflex  reaction  cannot  be  really  in- 
telligible to  the  physiologist  until  he  knows  its  aim."  ^^  No 
biologist  whose  mind  is  both  open  and  reasonably  pene- 
trating will  hesitate  to  accept  these  views  as  correct. 

What  I  wish  to  call  particular  attention  to  is  that  the 
last  sentence  quoted  calls  for  an  additional  clause:  "the  re- 
flex reaction  cannot  be  reall}^  intelligible  to  the  physiologist 
until  he  knows  its  aim,"  and  he  can  know  its  aim  only  hy 
considering  it  in  the  light  of  the  organism's  entire  complex 
of  normal  activities;  i.e.,  in  accordance  with  the  conception 
of  the  organism  as  a  whole. 

REFERENCE  INDEX 

1.  Sherrington  6       14.  Sherrington  136 

Q.  Sherrington  12       15.  Sherrington    205 

S.Sherrington  12       16.  Cannon,  W.  B.  ('16)   35 

4.  Sherrington  38       17.  Cannon,  W.  B.  ('16)   34 

5.  Sherrington  8       18.  Cannon,  W.  B.  ('14)   262 

6.  Donaldson 605       19.  Cannon,  W.  B.  ('16)   31 

T.Sherrington  16       20.  Cannon,  W.  B.  ('16)    34 

S.Sherrington  8       21.  Cannon,  W.  B.  ('16)   107 

Q.Sherrington  116       22.  Cannon,  W.  B.  ('16)   108 

10.  Sherrington  145  23.  Cannon,  W.  B.  ('16)   185 

11.  Sherrington  146  24.  Sherrington    114 

12.  Sherrington  151  25.  Sherrington    236 

13.  Sherrington  136 


Chapter  XXI 

IMPLICATIONS    OF    THP^.    TllOriSTIC    AND    SEG- 
MENTAL THEORIES  OF  NERVE  ACTION 

HAVING  now  carried  our  study  of  the  nei'vous  sys- 
tem far  enough  to  enforce  the  conception  that  the 
very  essence  of  this  system,  even  the  wholly  reflex  aspect  of 
it,  is  its  unifying,  its  integrating  office  for  the  organism,  it 
remains  to  see  what  the  elementalist  mode  of  reasoning  can 
do  with  the  system  in  this  aspect  of  it. 

As  in  the  case  of  internal  secretions,  we  can  get  at  this 
subject  in  no  way  better  than  tlirough  tlic  writings  of 
Jacques  Loeb.  Tliis  way  of  approach  will  be  seen  to  bo  tlic 
more  advantageous  when  Loeb's  long-continued  and  dis- 
tinguished experimental  studies  on  some  of  the  activities  of 
several  kinds  of  lower  animals  are  borne  in  mind. 

Neglect  of  the  Works  of  Slierrington  and  Cannon  hjj 

Jacques  Loch 

Despite  Loeb's  express  statement  in  the  preface  to  Thr 
Organism  as  a  Whole  that  the  volume  is  intended  to  be  a 
companion  to  The  Comparative  Fhijsiologif  of  the  Brain,  on 
which  account  a  discussion  of  the  central  nervous  system  is 
omitted  from  The  Organism  as  a  Whole,  the  omission  seems 
very  unfortunate.  Much  important  work  on  tlie  physi<)h)gy 
of  the  nervous  system,  especially  on  Its  integrating  function, 
has    been    done    since    The  Comparative  Thysiology  of  the 

185 


186  The  Unity  of  the  Orgamsm 

Brain  was  published,*  Sherrington's  realW  epochal  book 
being  especially  notable  among  these  later  productions. 
Some  of  the  most  fundamental  views  and  arguments  pre- 
sented by  Sherrington  and  others  are  irreconcilable  with 
conceptions  which  are  at  the  very  center  of  the  elementalist 
philosophy,  and  which  find  repeated  expression  in  Loeb's 
writings  on  the  nervous  system.  It  is,  consequently,  a  mat- 
ter of  surprise  and  disappointment  that  a  volume  devoted 
expressly  to  the  organism  as  a  whole  should  leave  almost 
unmentioned  the  very  organ-s^^stem  which  is  most  distinctive 
of  the  organism  thus  viewed,  especially  since  this  leaves  un- 
touched clearly  formulated  theoretical  issues  of  cardinal 
importance. 

Likewise  the  disposal  of  Cannon's  work,  upon  which  we 
have  drawn  so  largely  in  the  preceding  chapter,  by  five 
lines  in  The  Organism  as  a  Whole  ^  is  truly  astonishing ! 

Indeed,  so  striking  is  the  defectiveness  of  The  Organism 
as  a  Whole  in  this  respect  that  one  can  not  help  recognizing 
that  despite  its  author"'s  statement  in  the  preface  as  to  his 
intentions,  the  omission  amounts  to  an  evasion  even  though 
not  so  intended. 

However,  the  expressed  statement  by  Loeb  of  the  place  he 
wishes  the  Comparative  Physiology  of  the  Brain  to  hold 
in  his  interpretation  of  the  organism  has  great  usefulness 
to  us :  it  is  tantamount  to  an  assurance  that  the  views  set 
forth  in  the  Comparative  Physiology  have  undergone  no 
essential  change  because  of  the  discoveries  and  arguments 
produced  since  the  publication  of  that  work.  So  our  exam- 
ination of  Loeb's  position  on  this  important  matter  must 
concern  itself  largely  with  facts  and  views  contained  in 
the  Comparative  Physiology  of  the  Brain. 

*  The  copyright  of  Comparative  Physiology  of  the  Brain  bears  the 
date  1900,  while  The  Integrative  Action  of  the  Nervous  System,  by 
Sherrington,  than  which  few  more  important  books  on  the  system  have 
been  written,  was  copyrighted  in  1906. 


Implications  of  the  Theories  of  Nerve  Action      187 

The  Real  Importance  of  Loch's  Conception  of  the  Xervous 

Si/ stem 

First  of  all,  it  is  witli  genuine  satisfaction  that  I  recognize 
the  eminent  service  rendered  plivsiolocry  aiid  "-eneral  hioloffv 
by  this  author's  experimental  investigations  on  animal  ac- 
tivities. I  have  long  considered  that  one  of  physiology's 
foremost  achievements  is  the  clarification  of  the  conception 
that  nerve  phenomena,  the  most  characteristic  features  of 
which  are  response  to  stimuli  and  the  conduction  of  the 
effects  of  stimulation,  are  not  wholly  unique  attributes  of 
nerve  tissue,  but  are  elaborations  of  attributes  inherent  in 
all  protoplasm.  Perhaps  no  single  pliysiologist  has  con- 
tributed more  to  tliis  clarification  than  has  Loeb.  Par- 
ticularly important  has  been  his  demonstration  tliat  nerve 
centers  in  the  sense  of  ganglionic  masses  which  determine 
reflexes  are  not  fundamental  to  the  coordinated  and  adapted 
movements  of  animals;  that  reflexes  take  place  in  manv 
animals  normally  and  in  many  others  experimentally,  where 
no  such  centers  exist,  as  for  example  in  the  ascidian  after 
the  single  ganglionic  mass  is  cut  out. 

The  pertinacity  and  teclmical  skill  with  which  T.oeb  has 
followed  up  these  ideas  are  highh'  commendable.  By  focus- 
sing attention  on  the  fact  that  since  plants  have  no  nei-Aous 
system,  they  are,  in  so  far  as  they  exhibit  movement  induced 
by  stimuli,  illustrations  of  the  principle  of  protoplasmic 
response  in  actually  differentiated  organic  beings ;  and  In- 
carrying  the  same  conception  over  Into  the  animal  world  and 
demonstrating  that  many  activities  here  also  are  dependent 
upon  direct  protoplasnuc  response  and  not  necessarilv  on 
a  specially  differentiated  ganglionic  mass,  he  was  led  to 
formulate  the  tropism  theory.  This  theory,  as  we  shall 
show  presently,  has  a  significance  which  I^oeb  himself  seems 
not  to  have  fully  comprehended,  his  general  standpoint  be- 
ing unconductive  to  such  comprehension.     The  theorv  must. 


188  The  Unity  of  the  Orgamism 

consequently,  hold  a  large  place  in  our  effort  to  see  what  Loeb 
as  a  representative  of  the  elementalist  school  in  biology  is 
able  to  do  with  the  integrative  action  of  the  nervous  system. 
Let  it  be  understood  that  with  the  general  controversy 
which  has  gone  on  so  long  and  warmly  concerning  the  value 
of  the  tropism  theory  for  explaining  animal  behavior,  we  are 
little  if  at  all  concerned  in  this  discussion.  The  point  of 
cardinal  importance  for  us,  and  the  thing  about  the  theory 
which,  as  mentioned  above,  Loeb  and  his  followers  seem  not 
to  have  perceived  clearly,  is  the  fact  that  the  theory  is 
really  dependent  for  so  much  of  validity  as  it  has  on  the 
conception  of  the  organism  as  a  whole.  In  other  words,  the 
theory  is  in  essence  an  organismal  and  not  an  elementalistic 
theory.  Hence  it  results  that  since  Loeb  is  the  author  and 
chief  proponent  of  the  tropism  theory,  but  is  at  the  same 
time  an  extreme  elementalist,  his  general  position  as  an  in- 
terpreter of  animal  activities  and  the  nervous  system  con- 
tains fundamental  contradictions.  Indicating  these  con- 
tradictions and  exploring,  though  incompletely,  the  course 
along  which  the  truth  lies,  will  form  a  natural  conclusion  to 
our  study  of  the  integrative  office  of  the  nervous  system. 

The  Organismal  Character  of  Tropisms  Partly  Recognized 

By  Loeb 

1  first  call  attention  to  the  fact  that  Loeb  himself  has  seen, 
though  "as  in  a  glass  darkly,"  the  organismal  character  of 
the  tropism  theory.  Speaking  of  the  lack  of  a  sharp  boun- 
dary between  reflexes  and  instincts,  he  says  some  authors 
are  wont  to  speak  of  reflexes  when  the  responses  to  stimuli 
pertain  to  single  parts  or  organs,  but  they  "speak  of  in- 
stincts where  the  reactions  of  the  animal  as  a  whole  are 
involved  (as  is  the  case  in  tropisms)."  ^ 

And  the  same  idea  comes  out  still  more  definitely  in  a 
proposal  to  discard  the  term  reflex  as  applied  to  such  a  re- 


Implications  of  the  Theories  of  Nerve  Action      189 

sponse  as  the  clasping  act  by  the  arms  of  the  decerebrated 
male  frog,  and  substituting  tlie  term  tropism.  "It  is  better,'' 
he  says,  "to  call  them  tropisms  since  the  organism  as  a  whole 
is  involved."  ^ 

But  Loeb's  most  illuminating  recognition  of  the  orgaiiis- 
mal  character  of  tropisms  is  found  not  in  any  positive 
acknowledgements  but  in  his  statements  of  what  a  tropism 
consists  of  essentially.  This  he  has  told  us  many  times  and 
in  varied  phraseology  during  the  last  twenty-five  years.  The 
following  from  The  Organism  as  a  Whole  will  serve  well  as 
the  basis  of  our  consideration :  "Animals  possess  plioto- 
sensitive  elements  on  the  surface  of  their  bodies,  in  the  eyes, 
or  occasionally  also  in  epithelial  cells  of  their  skin.  These 
photosensitive  elements  are  arranged  symmetricallv  in  the 
body  and  through  nerves  are  connected  with  synnnetrical 
groups  of  muscles.  The  light  causes  chemical  changes  in 
the  eyes  (or  the  photosensitive  elements  of  the  skin).  The 
mass  of  photochemical  reaction  products  formed  in  the 
retina  (or  its  homologues)  influences  the  central  nei'Aous  sys- 
tem and  through  this  the  tension  or  energy  production  of 
the  muscles.  If  the  rate  of  photochemical  reaction  is  equal 
in  both  eyes  this  eff*ect  on  the  symmetrical  muscles  is  equal, 
and  the  muscles  of  both  sides  of  the  body  work  with  equal 
energy ;  as  a  consequence  the  animal  will  not  be  deviated 
from  the  direction  in  which  it  was  moving.  This  lia])pens 
when  the  axis  or  plane  of  symmetry  of  tlie  animal  goes 
through  the  source  of  light,  ])rovided  only  one  source  of 
light  be  present."  * 

Reduced  to  its  lowest  terms,  the  tropistic  mechanism  di- 
scribed  here  and  in  many  other  places  consists  of:  first, 
portions  of  the  body  surface  having  specific  irritability,  as 
receptor  organs,  these  being  usually  paired  and  usually 
symmetrical,  though  often  unsymmetrical;  second,  muscles 
as  effectors,  almost  always  paired  and  usually  s^^mmetrical ; 
third,    nerves,    variously    constituted,    as    conducting    ])atlis 


190  The  Unity  of  the  Organism 

connecting  more  or  less  directly  the  sensory  areas  with  the 
muscles.  In  a  word,  tropistic  movements  are  determined  by 
a  definite  number  of  definite  kinds  of  parts  or  organs,  defi- 
nitely arranged  with  reference  to  one  another  and  to  the 
whole ;  that  is,  upon  an  organized  body  called  a  living  being, 
or  briefly  an  organism.  A  morphological  entity  is  funda- 
mental to  the  tropism  conception.  Tropisms  are  explained, 
partly,  by  these  organizations.  Loeb  does  not  avoid,  in  fact 
at  times  appears  not  to  try  to  avoid,  recognizing  this :  "The 
irritable  structures  at  the  surface  of  the  body  and  the  ar- 
rangement of  the  muscles  determine  the  character  of  the 
reflex  act."  "The  expfanation  of  them  [tropisms]  depends 
first  upon  the  specific  irritabihty  of  certain  elements  of  the 
body-surface,  and,  second,  upon  the  relations  of  symmetry 
of  the  body."  ^ 

The  Organismal  Character  of  the  Segmental  Theory  of  the 

Nervous  System 

Another  province  in  which  Loeb  has  done  distinguished 
work  and  into  which  the  organism  as  such  constantly  ob- 
trudes itself  and  will  not  accept  relegation  to  a  secondary 
place,  is  that  of  the  coordinated  action  of  the  central  nerv- 
ous system  in  those  animals  in  which  that  system  consists 
of  an  axial  series  of  ganglia.  Here  again,  as  in  the  tropism 
theory,  the  basal  perception  underlying  the  segmental  theory 
of  nerve  physiology  is  the  truth  that  nei've  "centers"  as  the 
"seats"  of  particular  activities  and  functions  in  the  old  sense 
(in  the  sense,  that  is,  of  being  the  real  source  of  these 
activities)  do  not  exist.  According  to  the  segmental  theory 
the  real  unit  of  activity  of  an  animal  made  up  of  metameres 
or  segments  is  the  segment  itself,  the  nerve  ganglion  of  the 
segment  being  only  one  element  in  the  complex.  The  idea 
is  set  forth  with  fullness  and  perspicacity  in  the  chapters 
of  the  Physiology   of   the  Brain  dealing  with  worms   and 


Implications  of  the  Theories  of  Nerve  Action      191 

arthropods,  and  especially  in  the  one  on  the  segmental  the- 
ory in  vertebrates. 

The  starting-point  chosen  by  Loeb  for  the  treatment  is 
the  eartliworm.  It  is  generally  known  tliat  tliis  animal  is 
jointed,  or  segmented,  that  every  segment  has  a  pair  of  nerve 
ganglia  situated  underneath  the  intestine  on  the  floor  of 
the  body  wall,  and  that  the  pairs  of  ganglia  are  all  connected 
with  one  another  fore-and-aft,  so  that  the  whole  constitutes 
a  chain  of  ganglia  extending  the  entire  length  of  the  bod  v. 
In  addition  to  this  series  of  ventral  ganglia,  there  is  a  pair 
of  large  ganglia  above  the  gullet  at  the  extreme  front  end 
of  the  creature,  these  being  connected  with  the  ventral  nerve 
chain  b}'^  commissures.  These  dorsal  nei'^-e  masses  are  usu- 
ally called  the  supra-esophageal  ganglia,  but  are  often 
spoken  of  as  the  brain. 

On  the  basis  of  the  "center"  theory  of  the  nei'vous  sys- 
tem, it  was  very  natural  to  raise  the  query:  Is  the  earth- 
worm's brain  the  "seat",  the  primal  source  of  the  coordi- 
nated activities,  the  crawling  and  burrowing  which  make 
up  so  large  a  part  of  the  life  activities  of  these  crea- 
tures.'' Loeb's  own  formulation  of  the  question  is  good. 
"Does  coordinated  progressive  movement  in  which  all  the 
segments  of  the  body  participate,  depend  upon  the  brain.'*" 

The  first  attempt  to  answer  this  question  experimentally 
seems  to  have  been  made  by  Friedlander.  Having  made  a 
small  opening  in  the  side  of  the  body  of  a  living  worm  and 
cut  out  a  piece  of  the  nerve  cord  so  that  a  complete  hiatus 
in  the  cord  was  produced,  the  author  fully  expected,  he  tells 
us,  that  after  the  healing  of  the  wound  in  the  body  wall, 
which  took  place  very  quickly  and  completely,  the  parts  of 
the  body  in  front  of  and  behind  the  section  of  the  cord  would 
cither  behave  like  two  separated  individuals,  or  that  the 
front  part  would  draw  the  rear  part  passively  about.  But 
neither  result  followed.  "Animals  from  the  middle  of  which 
a  piece   of  the   central  nervous   system   is   wholly   wanting. 


192  The  Uiiity  of  the  Organism 

crawl  exactly  like  normal  animals  except  as  to  a  small  dif- 
ference to  be  noticed  later."  ^  The  wave  of  muscle  con- 
traction and  other  movements,  starting  at  the  anterior  end, 
passed  over  the  point  at  which  the  ganglionic  chain  was  in- 
terrupted and  on  to  the  hind  end  of  the  body  just  as  though 
there  were  no  such  interruption.  "The  movements  were 
coordinated  exactly  in  the  same  way  as  in  the  normal  ani- 
mal." 

These  experiments,  together  with  others  that  have  since 
been  performed  by  other  investigators,  overthrow,  as  Loeb 
savs,  "the  idea  that  coordination  in  these  animals  is  deter- 
mined  by  a  special  centre  of  coordination  which  is  located 
in  the  brain."  ^ 

.  But  if  the  brain  is  not  the  coordinating  center  and  does 
not  contain  it,  what  and  where  is  that  center?  The  answer  is 
there  is  no  such  center.  Coordination  is  not  a  central  matter 
at  all.  Rather  it  is  a  matter  of  the  working  together  of 
many  parts,  each  duly  balanced  off  with  many  others,  and 
properly  subordinated  to  the  whole  animal. 

The  details  of  the  truth  tlms  stated  in  general  terms  can 
be  illustrated  more  advantageously  by  referring  to  some  of 
Loeb's  own  observations  on  a  group  of  annelids  considerably 
higher  in  the  scale  of  life  than  the  earthworm.  I  refer  to 
the  pile  worms  familiar  to  many  persons  accumstomed  to 
marine  shore  fishing.  This  group  of  worms,  known  collec- 
tively as  nereid  annelids,  are  far  more  "heady"  than  earth- 
w^orms  in  that  the  useful  body  member  named  head  is  more 
definitely  set  off  from  the  rest  of  the  body  than  it  is  in  the 
earthworm,  and  is  outfitted  with  eye-spots,  touch  appen- 
dages, and  so  on,  wholly  lacking  in  the  earthworm.  Nereis 
is  much  more  highly  "cephalized"  than  is  the  earthworm,  so 
brain  centralization  would  be  expected  to  present  features 
here  not  occurring  in  the  more  lowly  worm.  When  nereids 
are  deprived  of  the  supra-esophageal  ganglia  their  behavior 
is  such  as  to  suggest  at  first  sight  support  for  the  concep- 


Implications  of  the  Theories  of  Nerve  Action      19U 

tion  that  the  brain  is  a  fundamentally  originative  and  coor- 
dinativc  member.  For  example,  an  individual  thus  depleted 
will  crawl  about  unceasiiifrly  on  the  sand  of  the  a(|uariuiii 
bottom,  without,  however,  showing  any  signs  of  burrowing 
into  the  sand,  though  tliis  operation  is  very  characteristic 
of  the  normal  animal.  Tliis  looks  as  if  the  brain  were  indeed 
the  seat  of  the  burrowing  instinct,  liut  not  so.  'J'Ik-  bur- 
rowing activity  can  be  induced  not  only  in  animals  deprived 
of  the  brain,  but  even  in  parts  of  animals  from  which  the 
heads  have  been  cut  away  altogether.  Loeb  placed  a  brain- 
less piece  of  a  Nereis  on  the  sand,  and  as  usual  it  remained 
quiet.  "I  then  gradually  covered  the  forward  end  with 
sand.  The  rest  of  the  animal  immediately  began  to  make 
the  typical  movements  which  the  animal  makes  in  forcing  its 
way  into  the  sand.  At  the  same  time  the  glands  began  to 
secrete  the  sticky  substance  which  cements  the  particles 
of  sand  tog-ether,  forming  the  wall  of  the  burrow-hole." 
"But,  why,"  Loeb  asks,  "does  the  Nereis  not  burrow  when 
deprived  of  its  brain  .'^  For  the  simple  reason  that  it  makes 
use  of  the  organs  of  the  mouth  in  burrowing,  and  these  are 
amputated  with  the  head.  Hence  it  is  the  loss  of  the  i)e- 
ripheral  head-organs  which  keeps  the  decapitated  Nereis  from 
burrowing,  and  not  the  loss  of  the  brain."  ^ 

Since  the  coordinated  activities  involved  in  burrowing  can 
be  accomplished  through  the  acting  together  of  various  su- 
])erficial  and  deeper  body  parts — sense  organs,  nuiscles,  and 
so  on — without  the  intermediation  of  the  brain,  these  pe- 
ripheral members  are  the  real  "seat"  of  such  activities.  The 
author  then  goes  on  to  inquire  what  ])art  the  brain  does  })lay 
in  the  creature's  normal  life.  "If  we  conq)arc,"  he  says, 
"the  conduct  of  a  Nereis  whose  brain  has  been  anq)utated 
with  that  of  a  normal  worm,  the  difference  seems  to  \w  of  the 
same  nature  as  that  between  an  insane  and  a  rational  human 
being.  .  .  .  The  peculiar  irritability  by  means  of  which  the 
Nereis  draws  its  head  back  and  moves  backward  out  of  tlie 


194  The  Unity  of  the  Organism 

tube  depends  upon  organs  which  are  located  in  the  forward 
end  of  the  body  and  whose  sensory  nerves  go  to  the  supra- 
esophageal  ganglion."  ^ 

One  office  of  the  Nereis  brain  is,  therefore,  to  make  a 
great  complex  of  activities  normal,  or  "of  the  same  nature" 
as  sane  activities  in  "rational  human  beings."  Not  the  ac- 
tivities themselves  but  their  normality  are  dependent  on  the 
brain.  This  distinction  is  undoubtedly  general,  but  it  is 
definite  so  far  as  it  goes.  Beyond  question  to  the  student 
who  should  compare  the  movements  of  the  normal  and  the 
brainless  Nereis  with  sufficient  care,  the  distinction  between 
the  two  would  be  as  positive,  as  indubitable,  and  as  definable 
as  that  between  almost  any  two  complexes  of  organic  phe- 
nomena. The  unsatisfactoriness  of  the  distinction  is  not  its 
vagueness  but  its  generalness.  It  is  sufficiently  definite 
though  not  sufficiently  analytic. 

Just  what  the  brain  does  to  normalize  the  activities ;  in 
other  words,  how  the  change  of  character  of  the  worm's 
behavior — its  "restlessness"  and  general  insane-like  conduct 
• — results  from  the  absence  of  the  brain,  is  not  known.  Loeb 
makes  a  suggestion  toward  an  explanation,  but  recognizes 
what  he  off*ers  as  nothing  more  than  a  suggestion ;  so  to  go 
into  it  is  not  worth  our  while. 

But  there  is  an  obvious  usefulness  of  the  ganglia  and 
whole  neural  chain  in  such  animals  aside  from  the  general 
one  of  normalizing  behavior.  It  serves  as  a  common  con- 
ducting and  relay  system  between  the  peripheral  sense  or- 
gans and  locomotor  muscles.  Loeb's  statement  of  the  matter 
is  concise  and  may  be  adequate:  "The  central  nervous  sys- 
tem does  not  control  response  to  stimulation ;  it  merely 
serves  as  a  conductor  from  the  point  of  stimulation  to  the 
muscle  through  which  weaker  stimuli  may  pass,  and  pass 
more  rapidly  than  would  be  possible  if  the  muscle  were  stim- 
ulated directly. 

"In   the  Annelids   each   ganglion   is    a   relay   station   for 


Implications  of  tlie  Theories  of  Nerve  Action      19/5 

the  sensory  and  motor  nerves  of  the  corrcspon(Hiiir  sciriiu'iit. 
If  the  head  exercises  a  stronger  influence  upon  tlie  hehavior 
of  the  animal  than  any  otlier  segment,  as  in  Nereis,  for  in- 
stance, I  beheve  it  is  due  to  the  fact  tliat  in  the  oral  end 
more  kinds  of  irritability  are  jiresent  and  more  peripheral 
organs  are  difif'erentiated  (sense  organs,  mouth,  etc.)  than  in 
the  other  segments."  ^'^ 

Loeb  has  done  good  service  in  bringing  togctlicr  the  evi- 
dence for,  and  emphasizing  the  truth  that,  the  segmental 
make-up  of  the  vertebrate  head,  so  long  and  so  ardently 
studied  by  anatomists  and  cmbryologists,  is  as  important  a 
matter  for  neural  physiology  as  it  is  for  mor|)hol()gy,  and 
that  as  a  matter  of  fact  the  vertebrate  spinal  cord  works 
more  or  less  on  the  segmental  principle,  and  in  this  respect 
falls  into  line  with  the  central  nerve  axis  of  annelids  and 
arthropods. 

The  first,  or  one  of  the  first,  investigators  to  recognize 
this  truth  was  Schrader,  in  his  studies  on  the  activities  of 
frogs  whose  brains  liad  been  wholly  or  partly  removed.  The 
frequently  quoted  and  later  fully  confirmed  statement  by 
this  investigator  is  of  prime  significance  for  the  general 
truth  of  what  we  might  call  fore-and-aft  neural  integration 
in  vertebrates.  "The  series  of  experiments  we  have  given 
teaches  us  that  the  central  nervous  system  of  the  frog  can  be 
divided  into  a  series  of  sections,  each  of  which  is  ca])able  of 
performing  an  independent  function.  It  brings  the  central 
nervous  system  of  the  frog  into  closer  relation  with  the  cen- 
tral nervous  system  of  the  lower  forms,  wliich  consist  of  a 
series  of  distinct  ganglia  that  are  connected  by  connnissures. 
It  speaks  against  the  absolute  monarchy  of  a  single  central 
apparatus  and  against  the  existence  of  dift'erent  kinds  of 
centers,  and  invites  us  to  seek  for  the  centralization  in  the 
many-sided  coupling  of  relatively   independent   stations.'' ^^ 

As  statements  much  at  variance  from  this  concerning  the 
"central"    function    and    power    of    difKerent    ])arts    of    the 


196  The  Unity  of  the  Organism 

frog's  brain  are  not  uncommon,  and  as  the  facts  are  spe- 
cially important  for  us,  it  is  desirable  to  consider  the  evi- 
dence a  little  more  in  detail.  To  this  end  the  following  from 
Holmes  may  be  taken  as  an  autlioritative  summing  up  of 
results  so  far  reached  in  this  field.  "A  frog  with  the  brain 
removed  as  far  back  as  the  medulla  is  still  capable  of  per- 
forming regular  leaps  and  swinnning  movements  of  the 
limbs.  When  thrown  on  its  back  it  rights  itself,  and  still 
performs  compensatory  motions  when  tilted  or  rotated. 
Breathing  is  normal,  and  the  animal  swallows  pieces  of  food 
that  are  placed  in  its  mouth."  ^^ 

If  the  removed  portion  of  the  brain  is  made  yet  larger, 
by  including  the  anterior  part  of  the  medulla,  the  animal  is 
still  capable  of  performing  a  long  series  of  regular  activi- 
ties. "Locomotion  is  effected  mainly  by  creeping,  but  the 
frog  is  nevertheless  capable  of  spring^ing  in  the  ordinary 
manner.  In  the  water  it  swims  by  alternate  movements  of 
the  limbs.  .  .  .  The  breathing  and  swallowing  reflexes  are 
still  normal,  but  tlie  croaking  reflex  is  no  longer  performed. 
The  reflex  of  snapping  at  food  is  not  destroyed.  ...  If  a 
piece  of  meat  is  rubbed  against  the  frog's  nose,  the  animal 
snaps  at  it  and  uses  the  fore  legs  to  stuff  it  into  the  mouth. 
The  same  reaction  may  be  brought  about  by  using  the  finger, 
but  after  the  finger  is  seized  and  it  is  found  that  the  object 
is  too  large  to  be  stuffed  into  the  mouth,  the  frog  begins  to 
reject  the  morsel,  and  uses  the  fore  legs  to  push  it  away. 
Truly  a  remarkable  combination  of  reflexes  !"^- 

And  further :  "There  is  no  center  for  coordinated  locomo- 
tion in  the  medulla.  Disturbances  of  locomotion  begin  with 
the  fore  limbs.  If  the  medulla  is  cut  across  at  the  tip  of 
the  calamus  scriptorius,  the  animal  sinks  on  its  breast,  and 
the  fore  limbs  are  for  a  considerable  time  helpless,  although 
the  hind  limbs  are  capable  of  performing  vigorous  coor- 
dinated movements.  The  reason  for  this  is  that  the  injur}^ 
lies  so  near  the  region  from  which  the  nerves  of  the  fore 


Implications  of   tlw   Theories   of  Xerir   Aetiou       1!)7 

limbs  arise  that  tlie  niovc'iiiciits  of  tli(S(^  mnnlKTs  arc  very  nat- 
urally affected."'"*  It  is  tlie  infcrdipeiulence  fore-and-aft 
of  reflex  activities  like  those  here  Indicated,  that  is,  of  tlie 
fore  limbs  acting  witli  the  mouth,  and  the  hind  linihs  with 
the  fore  limbs,  that  su^-i>-ested  to  Loch  the  idea  of  "cliain 
reHexes,"  a  conception  of  special  interest  from  our  stand- 
point. 

That  the  segmental  mode  of  functioning  of  the  vertebrate 
nerve  axis  is  not  limited  to  the  lower  portion  of  the  great 
vertebrate  phylum  is,  as  Loeb  points  out,  sho\v?i  bv  such 
investigations  as  those  of  Goltz,  and  of  (ioltz  and  l^wald  on 
the  dog.  These  are  given  so  fully  in  the  larger  text-books 
of  physiology  and  are  thereby  so  readily  accessible  to  all 
specially  interested  in  the  subject,  that  a  very  brief  presen- 
tation will  suffice. 

Two  main  groups  of  reflex  centers  in  the  ner\e  cord  are 
made  out,  the  cervico-thoracic  and  the  lumbo-sacral.  The 
first  group  contains  the  center  for  movements  of  the  an- 
terior limbs,  for  res]:)iratory  movements,  for  acceleration 
of  heart  action,  for  dilation  of  the  pupil  of  the  eye,  and  for 
several  other  activities  at  the  anterior  end  of  the  body.  The 
lumbo-sacral  group  contains  the  center  of  movements  for 
the  posterior  limbs,  for  control  of  the  anal  sphincter,  the 
activities  of  the  generative  organs,  and  several  other  jiarts. 

The  existence  of  such  centers  in  the  dog's  cord  was  proved 
by  section  of  the  cord  at  different  levels  under  the  most 
careful  operative  conditions.  Had  the  investigation  st()j)ped 
with  experiments  of  this  sort  the  facts  brought  out  might 
have  been  interpreted  as  confirmatory  of  the  "seat''  notion 
of  a  nerve  center.  When,  however,  it  was  discovered  that  a 
dofi-  from  which  most  of  the  cord  is  removed  nuiv  live  for 
many  months  in  a  good  state  of  health,  all  the  vegetative 
functions  being  carried  on  almost  typically,  it  became  clear 
that  the  various  centers  of  the  cord  are  not  the  seats  of  life 
activities    in    any    such    fundamental    sense    as    was    earlier 


198  The  Unity  of  the  Organism 

supposed. 

For  one  thing  the  results  seemed  to  show  that  the  sym- 
pathetic nervous  system  shares  with  the  central  system 
more  profoundly  than  had  been  known  before,  in  determin- 
ing life  processes. 

The  conclusions  which  flow  with  apparent  certainty  from 
the  observations  are  well  summed  up  in  the  following:  "Un- 
doubtedly all  such  activities  [of  visceral  life]  may  subsist 
and  function  in  a  comparatively  normal  fashion  after  re- 
moval of  all  spinal  influence.  The  office  of  the  spinal  sys- 
tem in  regard  to  the  visceral  life  seems  to  consist  in  en- 
dowing these  functions  with  greater  energy,  and  in  con- 
ferring greater  stability  and  more  solid  equilibrium  on  the 
general  constitution  of  the  animal."  ^^ 

The  reader  will  hardly  fail  to  recall,  on  reading  this  state- 
ment about  the  function  of  the  mammalian  spinal  cord, 
Loeb's  statement,  quoted  some  pages  back,  about  the  nor- 
malizing function  of  the  Nereis  brain.  "Stabilizing"  the 
dog's  function  means  much  the  same  as  "normalizing"  the 
annelid's  functions.  Nor  should  the  reader  neglect  to  notice 
that  he  might  substitute  the  word  "integratedness"  for 
"equilibrium"  in  the  quotation  without  change  in  the  es- 
sential meaning  of  the  sentence. 

Critique  of  the  Elementalist  Attempt  to  Interpret  Tropistic 
and  Segmental  Theories   of  the  Function   of   the 

Nervous  System 

We  now  return  to  the  central  point  of  the  present  sub- 
topic,  namely  that  of  what  a  genuine  bio-elementalist  is  able 
to  do  when  confronted  by  the  facts  in  possession  of  modern 
physiology  bearing  on  the  tropistic  and  segmental  theories 
of  the  nervous  system. 

First  we  are  compelled  by  the  evidence  to  recognize  the 
general  soundness  of  the  doctrines.      Second,  we  recognize 


Implications  of   the  Theories   of  Xenr   Action       ID!) 

tluit  probably  no  biologist  has  dcalf  witli  tlu-  coniij)! ioii 
more  coniprchcnsively  and  illuniiiiatiii<rly  than  Loch.  Third, 
wc  find  that  his  espousal  and  able  treatment  of  tlu-  tluorics 
has  led  him  into  })ositions  so  tlioroiitihly  or<rajiismal  in  both 
essence  and  expression  as  to  be  quite  irreconcilable  with  his 
own  more  general  elementalistic  philos()])hy. 

It  remains  to  point  out  more  speciticallv  than  \\i  liavr 
wherein  the  organismal  implications  of  his  teacliings  us  ex- 
emplified by  his  writings  constitute  a  refutation  of  the 
elementalistic  implications  of  his  teachings  as  exemplitie<l 
by  his  strivings  after  an  "ultimate  exjilanation"  of  organic 
phenomena  in  the  terms  of  physics  and  chemist i- v. 

The  essence  of  the  irreconcilability  of  the  two  positions 
may  be  put  into  a  general  form  thus:  in  innumerable  state- 
ments and  definitions  found  in  his  discussions  of  these  the- 
ories, Loeb  is  compelled  to  introduce  the  organism  either  as 
a  whole  or  in  considerable  portions,  as  a  causal  exphinatif)n 
of  particular  phenomena  with  which  he  deals;  and  the  com- 
pulsion to  such  introduction  makes  it  impossible  for  him  or 
any  one  else  to  dispense  with  the  causes  thus  introduced  by 
resolving  them  into  ultimate  elements  of  any  sort,  whither 
organic  or  inorganic. 

Illustration  and  justification  of  this  general  statement 
must  be  given.  The  following  typical  sentence  may  intro- 
duce the  discussion:  "The  irritable  structures  at  the  sur- 
face of  the  body,  and  the  arrangement  of  the  muscles,  de- 
termine the  character  of  the  reflex  act."  ''' 

Notice  what  it  is  that  "determines"  the  character  of  the 
act.  "Structures"  and  "muscles"  do  it,  tliese  being  "ar- 
ranged" so-and-so,  the  irritable  structures  dcfinitclv  on  the 
surface  of  the  body  and  the  nuiscles  within  the  body.  The 
point  needing  special  attention  i^  tliat  not  cliennfal  com- 
pounds or  even  living  substances  but  strueturcs  organs — 
and  these  arranged;  that  is,  entities  which  neither  exist  nor 
can  exist  except  through  the  agency  of  an   organism,  enter 


200  The  Unity  of  the  Organism 

into   the   definition. 

That  the  structures  referred  to  and  all  others  of  the  or- 
ganism are  composed  of  chemical  substances  and  nothing 
else,  is  beyond  question.  Why,  then,  does  the  author  not 
bring  into  the  definition  this  truth  about  the  ultimate  make- 
up of  the  structures.^  The  central  aim  of  his  researches 
on  animal  activities  being  what  they  are — "a  physico- 
chemical  analysis  of  behavior,"  is  it  not  surprising  that  he 
should  be  satisfied  with  even  a  working  description  or  defini- 
tion of  a  reflex  act  which  takes  no  notice  of  the  results  of 
such  analysis.^  If  the  understanding  of  complicated  life 
phenomena  consists  in  resolving  them  into  their  simple  ele- 
mentary components  (see  the  first  sentence  of  the  Physi- 
ology of  the  Brain) ^  then  a  definition  of  a  tropism  that 
would  contribute  largely  to  an  understanding  of  it  should 
not  be  content  with  such  proximate  constituents  as  "irri- 
table structures,"  "muscles,"  and  so  on,  but  should  go  right 
back  to  the  ultimate  physico-chemical  elements. 

The  typical  elementalist  answer  to  these  restrictive  criti- 
cisms is  well  known.  It  is,  substantially,  that  the  understand- 
ing and  hence  the  definition  of  tropism,  or  for  that  matter 
of  any  other  life  phenomenon,  is  final  or  ultimate  only  when 
expressed  in  physico-chemical  terms.  The  criticism  sug- 
gested concerning  the  mere  proximateness  and  hence  inade- 
quacy of  a  definition  that  uses  such  terms  as  "structures," 
"muscles"  and  "body  surface"  is  allowed  to  have  a  large 
measure  of  validity,  the  usage  being  justified  mainly,  it  is 
claimed,  on  practical  grounds.  The  morphological  concep- 
tions involved,  it  is  held,  are  so  strongly  intrenched  in  bio- 
logical terminology,  indeed  are  so  necessary  in  a  historic 
and  subsidiary  sense,  that  it  is  very  inconvenient,  if  indeed  it 
is  not  impossible,  to  dispense  with  them.  In  a  word,  the 
contention  is  that  while  morphology  and  general  physiology 
are  necessary,  their  necessity  is  secondary  or  subsidiary  to 
physics   and  chemistry.      But  what   should  be  seen  in   this 


Implications  of   the  Theories   of  \erve  Action       'U)] 

with  the  greatest  ])ossi})le  claiit  v  is  the  error,  the  fallacy,  not 
of  objective  fact  hut  of  reasoninij.  Stated  in  general  ttruis, 
the  fallacy  consists  in  the  tacit  assuni])ti()n  tliat  the  indis- 
j^ensahility  and  ade({iiacy  of  physico-chiiuical  elements  to 
exj)lain  organic  heliavior  are  of  a  sort  that  exclude  tin-  in 
dispensability  and  adequacy  of  "structures,"  as  sense  or- 
gans and  muscles,  from  the  explanation  of  the  same  ])he- 
n  omen  a. 

To  go  into  the  cpistcmological  and  logical  necessities 
involved  in  the  situation  with  whicli  we  are  liere  confronted 
is  entirely  beyond  the  scope  of  this  volume.  Ilowevtr,  in 
the  interest  not  so  much  of  Truth  in  the  abstract  as  of 
healthy,  wholesome,  useful  science,  biologists  will  have  to 
cease  employing  such  deprecatory  epithets  as  "anthropo- 
morphism," "metaphysics,"  "rhetoric"  and  the  liki-  to  gain 
for  themselves  a  sense  of  security  in  the  use  of  language  and 
reasoning  which  can  not  endure  for  an  lioui*  the  s(  ai-fhliirht 
of  really  careful  thinking  and  expression. 

To  focus  the  general  statement  of  the  fallacv  on  the  par- 
ticular matter  in  hand,  one  must  see  that  both  factually 
and  epistemologically  the  organs  and  other  morphological 
and  general  functional  elements,  or  factors  found  by  the 
analysis  of  the  organism,  are  "fundamental"  or  "ultimate" 
for  the  phenomena  to  be  explained  by  exactly  the  same  cri- 
teria that  the  physico-chemical  elements  are  fundamental 
or  ultimate. 

A  reflex  act  or  a  tr()])hisin  can  no  more  be  intelligibly 
expressed  or  understood  or  conceived  as  an  ol)')ectivi'  fact 
without  sense  organs,  muscles,  etc.,  than  without  physico- 
chemical  substances.  If  one  (piestions  the  truth  of  this  af- 
firmation let  him  test  the  matter  by  trying  to  express  a  re- 
flex act  in  the  terms  of  the  physico-chemical  elements  known 
from  analysis  to  be  "behind"  such  an  act.  To  In^gin  with,  he 
finds  it  necessary  to  fix  u])on  some  juirticnldv  reflex  act, 
scratch-reflex,  perhaps.     Such  })articular  act  nnist   be  taken 


20S  The  Unity  of  the  Orgamism 

as  the  starting  point,  otherwise  the  experimenter  will  be  com- 
mitted to  the  Idea  of  the  act  rather  than  the  act  itself. 
And  I  take  it  for  granted  that  no  truly  present-day  biologist 
is  willing,  even  though  he  may  not  be  able  to  justify  fully  his 
unwillingness,  to  commit  himself  to  that  ancient  position. 
Very  well  then,  the  scratch-reflex  is  going  to  be  expressed  in 
terms  of  the  oxygen,  nitrogen,  carbon,  phosphorus,  and  so 
on,  which  we  are  sure  are  "behind"  it.  Having  recognized 
the  necessity  of  starting  with  some  particular  reflex  act,  and 
having  settled  upon  the  scratch-reflex,  true  to  his  recogni- 
tion of  the  necessity  for  particularity  he  sees  he  must  go 
still  further  in  the  same  direction.  If  his  analysis  is  to  be 
exhaustive  and  his  expression  adequate  and  final,  it  will 
have  to  particularize  still  further  and  concern  itself  with 
the  reflex  of  some  particular  animal  species,  very  likely  even 
with  some  single  individual  or  small  group  of  individuals. 
A  vast  mass  of  evidence  makes  it  almost  certain  that  a 
dog's  scratch-reflex  is  different  from  a  cat's,  and  both  are 
different  from  an  ox's,  a  frog's',  and  so  on.  Suppose,  then, 
the  dog's  reflex  settled  upon,  and  defined  so  fully  that  it 
is  distinguished  from  every  other  reflex  whatever.  The  stu- 
dent is  now  ready  for  his  main  undertaking,  that  of  ex- 
pressing the  dog's  scratch-reflex  in  terms  of  the  physico- 
chemical  elements  to  which  the  reflex  is  reducible. 

The  next  step  is  to  examine  the  chemical  elements  con- 
cerned, as  these  are  treated  in  inorganic  chemistry,  for  the 
purpose  of  seeing  what  their  "terms"  are;  that  is,  what 
their  attributes  or  properties  are,  the  special  purpose  of  this 
examination  being  to  ascertain  what  terms,  i.e.,  what  at- 
tributes, answer  to,  or  correspond  to,  and  so  can  be  used  to 
express,  the  dog's  scratch-reflex.  The  outcome  of  this 
examination  is  unequivocal.  It  finds  no  terms,  no  attri- 
butes whatever,  which  by  themselves  suggest  even  remotely 
the  reflex  under  consideration. 

But  since  the  examiner  has  satisfied  himself  that  no  other 


Implications  of  the  Theories  of  Nerx^e  Action      5i()3 

simple  elements,  material  or  iininaterlal,  arc  "IkIiIiuI"  the 
refiex,  and  since  the  reflex  is  an  iiuhiliitable  reaHty,  tliere  is 
no  escape  from  tlie  conclusion  that  sometliing  other  tlian  the 
original  inorganic  simples  must  liave  intervened  hetweeii 
these  simples  and  the  reflex.  And  what  is  that  something? 
Docs  common  experience  and  connnon  sense  hesitate  with  its 
answer?  If  it  does  its  hesitancy  is  |)r()l)ahlv  from  surjjrise 
that  so  obvious  a  matter  should  Ik-  made  tlie  sul>/)i'ct  of 
serious  questioning.  "The  dog  is  what  lias  intervened  be- 
tween the  chemical  simples  and  the  reflex."  That  is  what 
common  experience  nmst  answer  and  will  unhesitatingly 
answer  once  it  recovers  from  its  surprise  at  being  (jues- 
tioned  on  a  subject  so  open  and  daylight  clear. 

But  then  science  comes  forward  with  its  criticism  of  this 
common-sense  answer.  There  is  no  gainsaying,  it  admits, 
the  truth  of  the  naive  answer  thus  given.  Ihit  this  answer, 
science  says,  is  a  mere  truism.  It  leaves  the  case  just  where 
it  was  before  science  began  its  analysis,  so  is  worthless  for 
scientific  purposes,  however  useful  it  may  be  for  ordinary 
purposes. 

This  rejoinder  by  science  nuist  be  looked  into  caicfully; 
otherwise  its  weakness  will  be  missed.  It  must  be  exacted 
of  science  that  she  show  more  explicitly  than  she  has  what 
she  means  by  explaining  the  dog's  scratch-reflex  by  refer- 
ring it  to  the  physico-chemical  elements  at  the  l)asis  of  the 
act.  Let  us,  we  must  insist,  hear  you  express  a  dog's 
scratch-reflex  in  the  terms  of  oxygen,  carbon,  et  cetera. 

That  such  expression  is  possible  is  freely  granted.  Hut 
how  can  it  be  done?  That  is  a  key  (piestion.  It  can  l>e 
done  in  one  and  only  one  way,  namely  by  adding  to  the 
attributes,  that  is,  to  the  "terms"  which  inorganic  chemistry 
recoirnizes  in  the  chemical  ileim-iits  concenied,  iust  those 
attributes  and  terms  which  the  dog's  scratch-reflex  requires 
in  order  that  the  elements  may  explain  the  reflex.  We  can 
say  that  besides  the  si>ccific  gravity,  combining  weight,  and 


204  The  Unity  of  the  Organism 

other  well-known  attributes  of  the  elements,  they  possess 
dog's-scratch-reflex  attributes.  But,  the  name  thus  sug- 
gested for  the  newly  discovered  attributes  being  cumber- 
some, we  may  devise  for  them  some  term  more  simple  and 
convenient — for  example,  do-sca-re-x,  doscarex. 

In  virtue,  then,  of  the  doscarecious  powers  of  oxygen, 
carbon,  and  so  forth  (the  fact  that  these  powers  can  only 
be  assigned  in  the  lump  and  not  distributively  to  the  several 
elements  should  not  be  lost  sight  of  though  it  may  be  neg- 
lected for  the  present  argument),  a  complete  physico-chem- 
ical explanation  of  the  phenomenon  under  consideration  is 
reached. 

Does  this  discussion  advance  the  interpretation,  the  under- 
standing of  biological  phenomena  beyond  the  familiar  sar- 
casm about  explaining  drug-induced  sleep  as  due  to  a  dor- 
mitive  principle  of  the  drug?  Yes,  I  maintain,  it  does, 
because  in  place  of  a  "principle"  attached  to  no  particular 
drug  but  to  any  sleep-producing  substance,  the  doscarecious 
powers  recognized  by  us  are  definitely  assigned  to  the  chem- 
ical elements  known  to  be  the  sole  constituents  of  organ- 
isms. The  new  powers  take  their  place  perfectly,  definitely 
and  positively  among  the  other  attributes  of  the  elements, 
the  assignment  being  based  on  the  solid  ground  of  analyses, 
laboratory  and  other,  made  through  years  of  scientific  re- 
search. But  an  exceedingly  important  point  which  comes 
in  sight  here  is  that  though  these  doscarecious  powers  are 
proved  to  be  real  ones,  they  are  latent  and  wholly  unknown 
to  inorganic  chemistry  for  tlie  reason  that  they  never  mani- 
fest themselves  under  any  other  combination  of  things  than 
just  that  which  in  its  totality  Zoology  has  named  dog.  A  dog, 
and  a  dog  only,  is  able  to  cause  oxygen,  carbon  and  the 
other  elements  to  reveal  these  particular  scratch-reflex 
powers.  The  dog  comes  in  as  a  sine  qua  non  to  the  pro- 
duction of,  and  hence  to  the  causal  explanation  of,  the 
particular    group    of    activities    under    consideration.      We 


Implications  of  the  Theories  of  Nerve    iction      205 

liave  no  evidence  that  the  chemical  elements  oiJcratliiL''  them- 
selves  can  actualize  their  own  latent  doscarecious  powers. 

The  reader  will  hardly  fail  to  connect  what  we  are  say- 
ing with  the  familiar  phenomena  of  the  assimilation  hv 
organisms  of  nutrient  suhstances.  All  our  aririiinciit  really 
does  to  the  usual  conce])tion  of  this  pheiiomiMoii  is  to  focus 
attention  upon  the  fundamental  im|)ortance  of  the  iml'n'idu/tl 
organism  as  a  factor,  as  a  cause,  of  the  chemical  transfor- 
mations wrought  in  the  nutrient  substances.  The  insnf 
ficiency  of  statement  of  the  assimilative,  oi-  anabolic,  or 
synthetic  aspect  of  the  metabolic  cycle  in  the  organism, 
lies  in  its  failure  to  bring  out  clearly  enouirh  the  Indubitable 
fact  that  the  final  results  are  innumerable  activities  and  sub- 
stances which  pertain  solely  to  the  living,  normal  indiridual 
— which  are  strictly  personal  and  private,  as  one  may  say. 

The  current  mode  of  expression  according  to  which  the 
assimilative  syntheses  take  place  in  the  organism  is  (juite 
misleading  in  that  it  permits  or  even  encourages  a  concep- 
tion that  the  syntheses  have  a  measure  of  detachment  from, 
and  independence  of,  the  life  of  the  organism  as  a  working 
unit,  which  as  a  matter  of  fact  they  do  not  have.  Again,  the 
usual  statement  that  the  syntheses  result  in  orga?iic  sub- 
stances of  more  complex,  higher  make-up  is  inadetjuate  in 
that  it  diverts  attention  from  the  fact  that  these  new  sub- 
stances belong  to,  are  fundamentally  ])ai"t  and  parcel  of  trie 
orxj-anism.  Thev  are  not  "any  old  organic  substances**  but 
are  exactly  those  substances  necessary  to  maintain  the  nor- 
mal  life  of  the  ])articular  individual  organism.  IKiui-  de- 
spite the  indubitable  fact  that  the  final  results  an-  reached 
by  w^ay  of  innumerable  |)urely  pliyslcal  and  chemical  ojH'ra- 
ations,  the  organism  itself,  acting  as  an  integer  no  niattir 
how  complex,  is  always  to  the  fore  as  a  controlling,  donn- 
nating  factor. 


206  The  Unity  of  the  Organism 

The  Bearing  of  This  Critique  on  "Analysis"  in  Biological 

Reasoning 

The  considerations  thus  briefly  set  forth  lead  to  certain 
still  more  general  ideas  of  the  utmost  importance.  The 
natural  entities  to  which  we  apply  the  descriptive  terms 
living  and  organic  are  specially  distinguished  by  the  chem- 
ical and  physical  syntheses  which  they  accomplish  by  virtue 
of  their  inherent  constitution.  So  as  concerns  the  most 
characteristic  of  these  syntheses,  especially  the  chemical 
ones,  they  are  to  a  very  considerable  and  fundamental  ex- 
tent definitive  of  the  organic  individual,  species,  genus,  and 
so  on,  of  taxonomic  biology.  This  is  equivalent  to  saying 
that  the  synthetic  operations  regarded  each  by  itself  ter- 
minate in  results  which  are  in  large  measure  unique  and  so 
unforeseeable  from  anything  we  know  about  the  original 
elements  as  such;  that  is,  before  they  have  actually  been 
subjected  to  the  particular  synthetic  transformations  under 
consideration. 

And  this  again  is  equivalent  to  saying  that  synthesis — 
transformatory  synthetic  processes  and  products— is  more 
distinctive  of  living  beings  than  are  analytic  products  and 
processes. 

Finally,  it  follows  from  these  facts  about  the  synthetic 
nature  of  organisms,  and  from  the  established  principles  of 
thought,  that  analysis  alone  is  incapable  of  intei'preting,  of 
understanding  organic  beings.  No  natural  object  which  in 
its  nature  is  more  distinctivel}^  synthetic  than  analytic  can 
be  understood  by  knowledge-processes  which  are  more  ana- 
lytic than  sjmthetic. 

Tliis  conclusion  goes  to  the  very  heart  of  the  elementalistic 
position,  and,  as  stated  in  the  discussion  on  internal  secre- 
tions, is  really  as  much  an  epistemological  as  a  biological 
problem. 

Reverting   again    to   Loeb's   writings,   the    conclusion    to 


Implications  of  ihc  Theories   of  Ner-r  Action      207 
which  WL'  are  forced    joins  issue  witli   the  very  oiK'ninf  sen- 

•.I  ^^ 

tence  of  tlie  Phi/sioloyi/  of  the  Brain,  as  we  have  ahM-ady  in- 
dicated. "The  understandinfr  of  c()ni|)licated  i>hen()niena," 
particularly  as  presented  by  orfranisms,  can  not  \ye  accom- 
plished tlirou"h  "an  analysis  by  which  they  are  resolved  into 
their  simple  elementary  components."  Unquestionably  an- 
alysis is  essential  to,  but  it  is  not  adequate  for,  full  under- 
standing of  the  phenomena. 

The  very  nature  of  organic  synthesis  and  of  hnojcledi/e- 
getting  precludes  the  possibility  of  attaining  the  hind  and 
degree  of  understanding  of  organisms  xchicJi  eU'mentalist 
biology  claims  to  have  attained  and  promises  to  attain. 

"A  com])lete  explanation  of  life  in  terms  of  physics  and 
chemistry"  is  impossible  for  the  sufficient  reason  that  physics 
and  chemistry  ^.9  such  do  not  contain  any  of  the  really  dis- 
tinctive  terms  of  life.  Those  terms  can  only  be  broufirht 
into  physics  and  chemistry  after  and  not  before  the  phe- 
nomena of  life  have  been  searchingly  scrutinized  ;  that  is,  an- 
alyzed and  found  to  involve  physics  and  chemistry.  The 
terms  of  life  are  in  the  original  data  on  the  phenomena  of 
life,  and  no  sort  of  analysis  can  possibly  make  this  other- 
wise. 

Our  position,  it  should  be  noticed,  touches  the  hackneyed 
controversy  over  vitalism  and  materialism  only  in  so  far  as 
the  course  of  reasoning  we  have  pursued  involves  the  recog- 
nition that  each  organism  (the  dog,  for  instance)  is  a 
natural  object  possessed  of  certain  causal  ])owers,  by  ex- 
actly the  same  logical  and  epistemological  criteria  that  any 
simple  chemical  element  or  chenucal  compound  is  a  natural 
object. 

Stated  in  a  brief  and  common-sense  way,  our  contention 
is  that  the  attributes  which  make  a  dog  a  living  body  an-  no 
less  natural  than  are  the  attributes  which  make  carbon  a 
chemical  body.  The  tiresome  and  meager-frnit*(l  contro- 
versy between  Materialists  and   N'italists  may  l)e  ch/iracter- 


208  The  Unity  of  the  Organism 

ized  as  due  to  the  fact  that  neither  party  has  taken  the 
trouble  to  estabhsh  clearly,  even  in  their  own  minds,  the 
meaning  of  the  word  natural.  As  a  consequence  of  this 
slipshodness  the  two  groups  agree  tacitly,  in  treating  the 
inorganic  and  the  organic  worlds  as  though  natural  does 
not  mean  the  same  thing  in  the  two  realms.  The  implica- 
tion is  that  if  the  inorganic  world,  for  instance,  be  held  to 
be  natural  by  both  parties,  for  the  Vitalists  the  living  realm 
is  largely  ^z^per-natural,  while  for  the  jNlaterialists  the  same 
realm  is  largely  iw/ra-natural. 

Theories  of  Animal  Behavior  in  Relation  to  the  Science  of 

Zoology 

This  somewhat  protracted  though  wofully  insufficient 
treatment  of  neural  integration  may  close  with  a  brief 
section  on  some  of  the  still  larger  biological  and  methodo- 
logical implications  of  the  conclusions  reached.  Special  at- 
tention is  called  to  the  fact  that  the  culminating  part  of 
our  argument  has  involved  data  and  fconceptions  which  are 
as  unequivocally  zoological,  morphological,  and  physiologi- 
cal, as  any  of  the  data  and  conceptions  are  unequivocally 
physical  and  chemical.  Physical  chemistry,  or  any  other 
aspect  of  inorganic  chemistry,  is  utterly  powerless,  so  far 
as  we  can  see,  to  discover  such  facts,  as  for  example,  that 
oxygen,  carbon,  nitrogen,  etc.,  possess  latent  "doscarecious" 
powers.  This  final  section  is  in  the  direction,  consequently, 
of  establishing  the  parity,  to  claim  the  least,  of  zoology, 
botany,  morphology,  and  general  physiology,  with  chemistry 
and  physics,  in  the  great  complex  group  of  biological 
science. 

We  may  first  allude  to  a  favorite  mode  of  expression  of 
materialistic  elementalists.  Whenever  fuller  analysis  has 
proved  some  group  of  animal  phenomena  not  hitherto  con- 
nected directly  with  jihysico-chemical  substances  and  forces. 


Implications  of   the   'lliconcs   of  Nerve  Action      ^Oi) 

to  be  in  reality  dependent  on  such  a^eneies,  this  sclior)!  !«, 
wont  to  remark  in  substance  that  investifration  lias  finallv 
"transferred"  the  phenonuiia  t'loiii  tlic  provinces  of  /ooln^v, 
in()r})hol()^y,  general  ])hvsioloi»v  and  the  othci*  sciences  of 
animal  life,  to  ])hy'Sics  and  chemistry.  Our  ar<rument  puts 
beyond  (]uestion  the  lof/ical  inadecpiacv  of  such  a  statement. 
Analysis  does  not  by  any  means  transfer  the  phenomena 
from  zoology,  etc.,  info  ]}liysics  and  chemist  ly.  Witlur 
analysis  nor  any  other  agency  can  any  more  take  the  study 
of  animal  phenomena  away  from  zoology  and  ])ut  it  into 
physics  and  cbcmistry  than  it  can  take  })read-making  away 
from  tlie  baker's  art  and  put  it  into  physics  and  cliemistry. 
The  chemist  may  undoubtedly  take  to  bread-making  and  find 
that  his  new  em])loyment  has  much  in  common  witli  his  old  ; 
but  in  so  far  as  he  really  succeeds  at  the  new,  he  is  more  a 
baker  than  a  chemist.  He  has  not  transferred  bread-making 
to  chemistry,  but  if  anything  has  done  just  the  reverse. 
What  analysis  actually  does  in  these  cases  is  to  extend  the 
bounds  of  physico-chemical  forces  and  laws  into  zoology, 
morphology,  etc.,  and  to  prove  that  if  zoological,  morpholog- 
ical and  physiological  undertakings  are  to  move  into  ever 
greater  fullness,  aid  from  physics  and  chemistry  is  indis- 
pensable. 

Thus  critical  examination  of  the  reasoning  of  elemen- 
talist  biology  reveals  the  logical  falhictj  in  any  sort  of  state- 
ment which  involves  the  assumption  that  the  older  sciences 
of  organic  beings,  like  taxonomic  botany  and  /oology,  geo- 
gra])hical  distribution,  mori)hology,  general  physiology  and 
so  on,  are  not  and  never  can  Ix'  relegated  to  j)laces  of  minor 
or  secondary  importance  in  biology.  Hut  it  is  the  praetieid 
liarmfidness  of  such  assumj)tions  rather  than  the  logical 
fallacies  underlying  them  which  cliietly  concern  us  in  this 
volume,  and  no  part  of  our  whole  subject  is  more  vitally 
aifected  by  such  harmfulness  than  this  of  the  InOiaNior,  even 
the  purely  tropistic  behavior,  of  animals.     The  whole  round 


210  The  Unity  of  the  Organism 

of  animal  biology — attitude  toward  research  problems  and 
undertakings,  valuations  and  interest  in  different  fields  of 
knowledge,  educational  aims  and  method — all  are  affected. 

In  the  light,  for  instance,  of  such  a  complex  of  animal 
behavior  as  that  presented  b}^  the  northern  fur  seal,  its 
mating,  breeding,  migrating  and  other  habits,  the  conten- 
tion that  the  myriads  of  complicated  phenomena  which  go 
to  make  up  animal  life  can  be  understood  by  converting 
zoology  into  a  laboratory  and  experimental  science,  to  the 
end  of  analyzing  the  phenomena  into  their  "simple  elemen- 
tary components,"  is  so  ludicrous  as  hardly  to  need  argu- 
mentative refutation.  Indeed,  it  seems  as  though  persons 
obsessed  by  a  theory  to  the  extent  of  being  impervious  to 
the  ludicrousness  of  the  contention,  are  likely  to  be  also 
impervious  to  the  true  reasoning  involved  in  it.  It  may,  I 
think,  be  assumed  that  so  much  of  zoology  as  has  formed 
this  remarkable  conception  of  itself  will  before  long  drop 
into  the  background  by  scientific  gravitation  despite  its 
present  great  vogue. 

One  of  the  leading  motives,  consequently,  of  this  con- 
structive part  of  my  entei'prise  is  to  establish  the  essen- 
tiality of  general  zoology  and  its  time-sanctioned  depart- 
ments on  so  solid  a  basis  of  philosophic  reasoning  that  the 
necessary  methodology  of  the  regenerated  science  of  the 
future  will  be  clearly  seen  in  broad  outline. 

If  the  considerations  inadequately  presented  in  these  last 
pages  and  in  other  parts  of  this  volume  once  get  secure 
lodgment  in  biological  thought  it  will  become  manifest  that 
the  "behavior"  of  any  animal  species  (as  of  the  fur  seal, 
to  take  at  random  any  one  of  thousands  of  species  that 
would  illustrate  the  point  quite  as  well)  can  mean  nothing 
less  to  a  really  scientific  biology  than  the  whole  series  of 
activities  of  at  least  one  individual  animal  from  its  birth  to 
its  natural  death.  Consequently  an  "understanding  of  the 
complicated  phenomena"  thus  presented  can  not  be  secured 


Implications  of  the  Theories  of  Xcrvc  Aciivn      211 

by  any  amount  whatever  of  analysis,  but  only  through  an 
endless  series  of  never-divorced  analyses  and  svntheses,  tills 
series  running  on  tlirough  years  of  effort  by  scores  of 
investigators  in  the  field  (on  the  Pribilof  Islands,  on  the 
Behring  Sea  and  far  down  into  the  North  Pacific  Ocean) 
and  in  the  laboratory,  by  general  zoologists,  mannnalogists, 
anatomists,  embryologists,  physiologists,  comparative  psy- 
chologists,  bio-chemists    and   ])hysical    chemists. 

How,   let   one  ask   himself,   would   the   resolving  of   a    fur 
seal's  behavior  into   reflexes   as   the  simple  elementarv   com- 
ponents of  that  behavior,  contribute  to  an  understanding  of 
the  annual  oceanic  migration  of  the  animal  or  the  fighting 
of  the  males  for  the  females,  if  that  analysis  accom})lished 
nothing  beyond  proving  a  certain  measure  of  identitv   be- 
tween the  reflexes   of  a  fur  seal  and,  for  instance,  those  of 
an   earthworm?      Or   how   would    the    understandin""   sou'dit 
be  enhanced  by  carrying  the  analysis  to  a  still  deeper  level 
— to  the  physico-chemical  level — and  discovering  just  how 
oxygen,  or  some  particular  protcid  substance,  participates 
in  the  reflex?     We  must  never  lose  sight  of  what  the  biolo- 
gist's task  is  as  regards  understanding.     It  is  to  investigate 
for   the  purpose   of   understanding  the   facts   presented   !)v 
living  beings  in  nature.     The  behavior  of  the  fur  seal  as  the 
animal  lives  its  normal  life  is  what  is  to  be  studied   in   tlu" 
case  chosen,  and   one  of  the  most  characteristic   things   in 
this  particular  behavior  is  a  yearly  journey  of  several  thou- 
sand miles  throuijh  the  Pacific  Ocean.      That  is  one  of  the 
phenomena  to  be  studied  and  understood,  and  no  amount  of 
analysis    resulting   in   discoveries   which   do    not    apply    par- 
ticularly  to   that   particular   phenomenon    can    be   admitted 
as  an  explanation  of  that  phenomenon.     As  com])ared  with 
any   of  the   other  natural   sciences,  biology   is    prei'ininently 
the  science  of  iniliridnals — or  natiii'al   objects  wliich  though 
alike    in    iiuiumerable    attributes    are    unlike    in    innumeral)Ie 
other  attributes,  and  in   no  as])ect  of  organisms  do  the  dif- 


212  The  Unity  of  the  Organism 

ferentials  which  make  individuals,  species,  genera  and  so 
on  come  out  so  importantly  as  in  behavior. 

The  evidence  being  now  overwhelming  that  all  organic 
phenomena,  including  beliavior,  are  dependent  upon  physico- 
chemical  substances  and  forces,  one  of  the  most  pressing  ques- 
tions of  procedure  in  biological  research  is  that  of  bringing 
the  older  and,  generally,  less  exact  natural  history  aspects  of 
the  science  into  closer,  more  vital  cooperation  with  its  newer 
experimental  and  more  quantitativel}^  exact  aspects.  Specif- 
ically stated,  work  of  the  type  long  prosecuted  by  explor- 
ing expeditions,  botanical  and  zoological  gardens,  museums, 
botanical,  zoological  and  biological  societies,  and  govern- 
ment biological  surveys ;  and  that  of  laboratories  in  the 
strict  modern  sense,  the  morphological,  physiological,  and 
bio-chemical  laboratories,  must  join  hands  more  closely  and 
effectively  than  they  have  heretofore  to  insure  continued 
progress  in  the  organic  sciences.  Several  movements  of  the 
day  in  biology  could  be  mentioned  whose  meaning,  viewed 
from  our  standpoint,  can  hardly  be  mistaken.  Perhaps  the 
most  conspicuous  of  these  is  that  congeries  of  research  ac- 
tivities known  as  ecology.  In  spite  of  frequent  deprecia- 
tive  comments  about  ecology,  especially  because  of  its  in- 
definiteness  as  to  both  content  and  delimitation,  it  has  the 
merit — from  our  standpoint  the  very  great  merit — of  facing 
organic  nature  as  it  actually  is,  that  is,  of  having  for  its 
subject  matter  the  modes  of  life  of  organisms  as  nature 
presents  them,  and  hence  of  recognizing  the  laboratory  as 
an  agency,  but  only  as  one  among  other  agencies,  for  deal- 
ing with  its  subject.  As  to  method,  while  ecology  recognizes 
the  indispensability  of  the  laboratory  and  experimentation 
in  the  narrow  sense,  it  refuses  to  let  such  experimentation 
usurp  the  whole  of  its  interest  and  effort. 

So  our  study  of  the  organism's  integratedness  as  exem- 
plified by  its  activities,  that  is,  by  its  behavior,  and  by  the 
mechanism   through  which    these   activities    are   carried   on. 


Implications  of  the  Theories  of  Nerve  Action      J2l;J 

leads  to  tlic  soniL'what  uiifxpcctcd  though  eiitirelv  natural 
result  wliicli  may  ))C'  suiuiuarily  stated  tlius:  To  ^ain  un- 
derstanding of  the  beliavior  of  living  beings  is  admitted  bv 
everybody  to  be  the  chief  reason  for  investigating  sucli  ac- 
tivities. Due  consideration  of  the  nature  of  the  activities 
and  of  the  nature  of  understanding  makes  it  certain  tliat  the 
])henoniena  themselves  are  lilghly  Integrative  and  integrated, 
or  synthetic,  and  that  understanding  of  tliem  de|)ends  as 
much  on  synthetic  knowledge-getting  as  on  analytic  knowl- 
edge-getting. Perception  of  this  last  truth  necessitates, 
again,  a  sort  of  synthesis,  or  integration,  of  the  numerous 
research  agencies. 


REFERENCE  INDEX 


1.  Loeb  ('16)    285 

2.  Loeb  ('12)    70 

3.  Loeb  ('16)    284 

4.  Loeb  ('16)    257 

5.  Loeb  ('02)    7 

6.  Friedlander    363 

7.  Loeb  ('02)    85 

8.  Loeb  ('02)    91 


9.  Loeb   ('02)    94 

10.  Loeb    ('02)    101 

11.  Schrader    177 

12.  Holmes    ('06)    313 

13.  Holmes    ('06)     315 

14.  Luciani    iii,  355 

15.  Loeb   ('02)    5 


Chapter  XXII 

PSYCHIC    INTEGRATION 

Preliminary  Remarks 

(a)     Absolute  Discrimination  Between  Reflex  and  Psychical 

Phenomena  Not  Necessary 

T  T  will  be  recalled  that  in  our  discussion  of  neural  inte- 
•*-  gration  we  limited  ourselves  strictly  to  those  manifesta- 
tions and  activities  of  organisms  which  are,  so  far  as  ob- 
servation can  determine,  strictly  reflex,  that  is,  show  no 
evidence  of  intelligence  and  volition,  or  even  necessarily  of 
instinct.  What  we  have  to  do  next  is  to  consider  the  unity, 
the  integratedness  of  the  animal  organism  as  manifested  in 
the  vast  array  of  its  activities  which  by  universal  consent 
are  designated  by  such  terms  as  instinctive,  emotional,  vo- 
litional, conscious  and  intelligent. 

Be  it  noted  that  this  aim  will  not  exact  of  us,  any  more 
than  did  the  last,  a  sharp  delimitation  between  reflex  or 
purely  mechanical  acts  and  psychical  or  conscious  acts. 
Just  as  in  the  former  discussion  we  were  concerned  with  the 
integrative  character  of  those  acts  which  are  indubitably 
reflex,  so  here  our  object  is  to  study  the  integrative  or  syn- 
thetic character  of  those  acts  which  are  indubitably  psychi- 
cal. We  shall  now  be  dealing  with  acts  which  have  unques- 
tionable psychical  attributes,  that  is,  show  something  of 
individual  plasticity  and  something  of  correspondence  to 
individual  needs ;  which  are,  in  other  words,  to  some  degree 
individually  determined  to  meet  individual  requirements 
either  of  external  or  internal  imposition  and  intention. 

214 


Psycliic   Integration  J815 

(h)     The  Organism  an  Origimd  Datum  in  All  J'rohltms  of 

Psychic    Life 

Another  preliminary  remark  of  liigJi  importance  concerns 
the  question  of  what,  precisely,  it  is  with  which  we  have  to 
do — of  what  we  start  from  and  what  is  ever  in  sirrht,  in  the 
discussion.  Our  fidelity  to  the  organism,  iivin<r  in  its  natural 
setting,  as  the  foremost  ohjective  reality  in  this  treatise, 
prevents  us  ah  initio  from  heing  satisfied  with  a  JJodv,  and 
a  Mind  or  Soul,  as  these  have  figured  from  time  innnemorial 
in  discourse  about  the  higher  animals,  particularly  ahout 
man. 

If  in  all  the  world  there  is  such  a  thing  as  objective 
truth,  what  we  start  with  and  have  ever  to  deal  with  in 
studying  psychic  phenomena,  just  as  in  studying  all  other 
phenomena -of  animals,  are  individual  objects  or  l)o(lies  of 
very  particular  construction  and  activity.  And  by  no  })os- 
sibility  can  consistent  thought  and  statement  avoid  acknowl- 
edging that  that  vast  assemblage  of  acts  and  other  manifes- 
tations which  are  called  psychical  are  yet  only  })art  and 
parcel  of  the  still  vaster  assemblage  of  acts  and  manifesta- 
tions presented  by  the  very  same  living  objects,  that  is,  by  or- 
ganisms. Our  occupation  will  be  basally  with  (in  object,  some 
particular  organism,  having  innumerable  attiM})utes,  which 
being  classified  fall  rather  roughly  into  two  great  groups, 
one  of  which  we  name  physical  or  material  and  tlu-  other 
psychical  or  spiritual.  For  short,  the  physical  or  material 
group  is  called  the  Body,  while  the  j)sychical  or  spiritual 
group  is  called  the  Soul  or  ]\rind. 

Our  discussion,  then,  will  luvci-  lose  sight  of  tlu'  fact  that 
the  acts  with  which  we  deal  arc-  acts  of  the  organism  and  not 
of  any  of  its  parts  merely,  wlutjui-  IIksi-  be  concei\i(i  as 
material  or  psychical.  No  matter  how  far  particular  acts 
may  be  dependent  upon,  and  so  explicable  by.  particular 
parts,  this  dependence  can  not   in  reality  i)e  the  whole  story. 


216  The  Unity  of  the  Organism 

for  the  sufficient  reason  that  the  parts  are,  finally,  non- 
existent except  as  derivatives  of  and  dependencies  upon  the 
organism,  as  our  whole  treatise  has  abundantly  shown. 

Discussion  of  the  Nervous  System,  the  Brain,  the  Cerebral 
Cortex,  Neurones,  Reflexes,  the  Senses,  Responses,  Emo- 
tions, Consciousness,  Will,  Reason,  and  so  on,  as  though  any 
of  these  are  now  or  ever  have  been  or  ever  will  be  independent 
entities,  or  things  to  which  the  organism  is  subordinate,  is 
from  our  standpoint  one  of  the  deep  inadequacies  and  mis- 
fortunes of  much  biological  and  psychological  thinking.  To 
the  whole  attitude  of  the  zoological  naturalist,  who  by  na- 
tive endowment  and  by  training  is  imbued  with  the  spirit 
of  his  motto  "neglect  nothing"  in  the  study  of  animals,  this 
habit  of  the  special  sciences  of  animal  life  is  intolerable. 

The  ancient  problem  of  the  relation  of  "Mind"  to  "Body" 
is  one  of  those  problems  which  run  on  endlessly  in  discussion 
simply  because  the  partisans  of  one  theory  or  another  never 
know  exactly  what  they  are  discussing — never  know  just 
where  they  start  from,  in  what  direction  they  are  going,  or 
what  the  end  would  be  like  if  they  reached  it. 

Lest  this  statement  be  taken  as  foreshadowing  both  a 
right  statement  and  a  "final  solution"  of  the  problem,  I 
affirm  very  positively  that  it  foreshadows  nothing  of  the  sort. 
All  I  hope  to  do  is  to  add  considerably  to  a  clear  statement 
of  the  problem,  to  add  a  little  to  our  comprehension  of 
whither  we  are  going,  and  to  contribute  a  bit  to  the  "final 
solution,"  whatever  that  may  mean. 

The  real  problem  of  psychic  integration  formulates  itself, 
for  us,  in  a  two-parted  way:  Given  any  particular  act  or 
action-system*  which  is  unquestionably  psychical,  (1)  how 
many  and  what  parts  of  the  organism  are  essentially  in- 
volved in  it?  and  (2)  does  the  act  or  action-system  bear  such 

*  This  phrase  I  borrow  from  Jennings  {Behavior  of  the  Lower  Organ- 
isms, p.  107)  and  mention  incidentally  here  that  we  shall  find  it  ex- 
tremely useful  later  on. 


Psychic   I ntcy ration  217 

relation  to  otlicr  acts  or  aciion-systcnis  and  to  iiialcrial 
parts  of  tlie  or^ranisin  as  to  warrant  the  ascription  to  these 
acts  of  causal  intluence  on  other  acts  and  on  tlic  materini 
parts? 

(c)   Provisional  Classification  of  Psychical  Facts 

A  third  and  final  introductory  remark  touches  on  tlu- 
question  of  what  shall  he  recognized  as  contained  in  the 
orp-anisni's  system  of  psychic  attrihutes.  FoUowinir  our 
regular  custom  of  beginning  with  the  ])lienomenon  under  con- 
templation at  its  fullest,  most  indubitable  expression,  we 
shall  not  go  far  amiss  if  we  accei:)t  the  time-honored  trium- 
yirate  of  feeling,  will,  and  intellect  as  the  most  obyious  suli- 
groups  of  highest  psychic  attributes ;  for  only  a  hopelessly 
sophisticated  })hilos()])hy  and  i)sych()l()gy  can  hesitate  to 
acknowledge  that  every  full-grown,  nonnal,  ciyihzed  human 
organism,  at  least,  is  at  once  a  sort  of  reservoir  of  feeling, 
sentiment,  and  emotion ;  a  dynamo  of  resolution  and  exe- 
cution;  and  a  granary  of  intelligence  and  reason.  (See,  for 
example,  Thinhing,  Feeling,   Doing,   by   K.   W.   Scripture.) 

Perhaps  the  only  thing  that  needs  saying  al)out  these 
sub-systems  of  mind  is  that  our  general  stan(lj)oint  aligns 
us  squarely  with  the  tendency  in  present-day  psychology  to 
accept  them  for  what  they  actually  are,  striying  to  become 
acquainted  with  them  and  to  assess  their  importance  on  this 
basis.  To  ascertain  first  of  all  the  facts  on  the  psychic  side 
of  the  hving  animal,  then  next  to  interpret,  to  correlate,  to 
explain  these  facts,  are  cardinal  principles  of  procedure  in 
our  enterprise.  For  one  thing,  as  an  evolutional  zoologist 
of  many  years'  practice  in  speculating  on  how  animal  parts 
originated  (even  those  of  almost  infinite  siinj)li(ity  as  com- 
pared  with  the  mind  of  man),  I  am  too  familiar  with  the 
limitations  and  ])itfalls  of  the  genetic  method  to  Ik.-  be- 
guiled into  making  souje  one  theory   of  the  origin   of  mind 


^1^  The  Unity  of  the  Organism 

the  corner-stone  of  my  interpretation.* 

Furthermore,  as  a  naturalist  faithful  to  the  mandate  "neg- 
lect notliing,"  I  am  in  full  accord  with  psychology's  aban- 
donment of  the  earlier  supposition  that  a  leading  aim  of 
psychology  must  be  to  prove  that  some  one  psychic  province 
is  all-dominant,  the  others  being  merely  secondary  and  trib- 
utary. 

Thus  all  forms  of  the  theory  that  the  psychic  empire  is 
at  bottom  Intellect,  the  heaven-ordained  monarch  of  which 
is  Unconditioned  Ideation,  are  incompatible  with  our  stand- 
point. Something  of  the  weight  and  variety  of  authorita- 
tive sanction  which  are  pitted  against  us  here  is  indicated 
by  such  names  as  Descartes,  Locke,  Leibnitz,  Hume,  Kant, 
and  Herbart  of  the  proximate  past,  and  Wundt,  Royce, 
Howison  and  Bradley  of  the  immediate  past. 

And  theories  like  those  of  Fichte,  Schopenhauer,  Hart- 
mann,  Nietzsche,  and  divers  pale,  fitful  present-day  lights, 
which  would  accord  to  Will  hegemony  over  the  entire  psychic 
realm,  are  still  less  tolerable  to  us. 

Feeling  has  won  its  rightful  place  in  psychology  so  recently 
that  there  seems  little  danger  of  its  pushing  its  claims  to 
position  and  power  beyond  reason.  Except  perhaps  in  the 
form  of  sensationalism,  or  the  theory  that  all  knowledge  ac- 
tually does  originate  in  sensations,  psychology  of  the  west- 
ern world  appears  never  to  have  attempted  seriously  the 
deification  f  of  Feeling  as  it  has  of  Reason  and  Will.     When 

*  I  count  it  as  one  of  the  pieces  of  good  luck  in  my  scientific  career 
that,  through  no  merit  of  my  own,  a  technical  memoir  of  mine  containing 
an  elaborate  theory  of  the  origin  of  the  vertebral  column  has  lain  in 
editorial  keeping  unpublished  for  a  decade  and  a  half. 

t  It  is  possible,  as  my  friend  Professor  G.  M.  Stratton  suggests  to  me, 
that  the  German  philosopher  Fr.  H.  Jacobi,  came  nearer  doing  this  than 
any  one  else.  His  teachings  gave,  however,  a  definite  place  to  positive 
knowledge,  so  that,  according  to  Hoffding,  his  faith  and  his  knowledge 
constituted  two  distinct  philosophies.  What  he  wrote  probably  does  not, 
consequently,  contradict  the  statement  in  the  text.  Jacobi  seems  not  to 
have  exerted  much  influence  on  the  main  current  of  German  philosophy 
and  life. 


Psychic  Integration  219 

Thomas  Hobbcs  Identified  Imatrlnution  willi  fancy,  ^'ori^r- 
inal  fancy"  with  sense,  and  "decaying  sense"  witli  nuinory, 
and  held  sense  to  bo  caused  by  "so  many  several  motions  of 
the  matter,  by  which  it  presseth  our  organs  diversely,"  and 
when  he  defended  tlie  general  (h)ctrine  that  "there  is  no 
conception  in  a  man's  nn'nd  wliich  hatli  not  at  fir.^t,  totally 
or  by  parts,  been  begotten  upon  tiie  organs  of  sense,"  ^  he 
did  indeed  blaze  a  trail  wliich  might  easily  lead  to  an  over- 
exaltation  of  the  sensuous  and  emotional  side  of  life.  Hut 
the  eminently  practical  character  of  Hobbes'  undertakiuL^ 
being  remembered  (he  was  writing  not  for  the  love  of  specu- 
lation but  to  save  his  country  from  political  chaos  and  mis- 
ery begotten  as  he  believed  from  false  theories  and  impossible 
desires  and  ambitions)  one  may  expect  to  find  in  him  ele- 
ments of  steadiness  and  restraint  which  would  make  for  safetv 
in  speculation. 

One  such  element  is  clearly  seen  in  his  opinion  that  he  who 
is  "born  a  man"  and  lives  "with  the  use  of  his  five  senses" 
has  all  the  native  equipment  necessary  to  realize  the  best  in 
him  both  for  himself  and  for  his  country.-  The  necessity 
of  being  "bora  a  man"  is  the  point  to  be  especially  noticed. 
That,  with  all  it  may  imply,  is  on  a  par  with  the  necessity 
of  using  the  five  senses.  So  whatever  of  scientific  hohl)y- 
riding  under  such  captions  as  "sensationalism,"  "empiric- 
ism," "associationalism,"  may  have  followed  in  the  wake  of 
Hobbes'  writings,  Hobbes  himself,  I  am  quite  sure,  was  at 
heart  a  genuine  organismalist  and  is  entitled  to  high  es- 
teem as  one  of  the  very  first  moderns  to  speak  strongly  for 
the  inq)ortance  of  the  body  generally  to  the  psychic  life  of 
man.     Listen  to  this  : 

"Natural  sense  and  imagination  are  not  subject  to  al^- 
surdity.  Nature  itself  cannot  err;  and  as  men  abound  in 
copiousness  of  language,  so  they  become  more  \\  isr,  or  more 
mad  than  ordinary."  "Between  true  science  and  errotuous 
doctrines,  ignorance  is  in  the  middle."  ^ 


220  The  Unity  of  the  Organism 

Having  completed  a  recoiinoissance  of  the  field  in  which 
we  are  to  work,  of  its  expanse,  its  contents,  and  its  main 
subdivisions,  we  are  prepared  to  take  up  the  task  proper. 
Approaching  it  from  our  standpoint,  one  naturally  sur- 
mises that  between  the  organism's  neural  unity  as  manifested 
by  its  reflexes  studied  in  the  last  chapter,  and  its  psychical 
unity  known  to  psycliology  and  to  be  considered  presently, 
a  vital  unity  of  still  higher  order  exists.  By  unity  I  mean  a 
unity  so  intimate,  so  reciprocating,  so  mutually  constitutive 
that  the  term  parallelisnfi,  with  the  meaning  given  it  in  much 
of  recent  psychology,  is  wofully  inadequate  for  it.  Such  a 
unity,  if  it  exists,  must  be  sought  by  inspecting  the  entire 
gamut  of  psychic  life,  from  the  simplest  responses  to  stim- 
uli on  up  through  simple  reflex  responses,  the  tropisms,  the 
primal  aff*ective  and  emotive  responses,  through  the  perceiv- 
ing, the  imagining,  the  conceiving,  the  reasoning  operations, 
to  the  very  highest   constructive  human  mental  achievings. 

Likeness   Between   Tropistic   tmd  Higher  Psychic   Activity 

An  important  move,  starting  from  the  highest  phase  of 
rational  mind,  has  been  made  toward  recognizing  the  nature 
of  this  unity.  This  move  is  the  more  significant  for  our 
enterprise  in  that  the  investigator  who  has  made  it  is  neither 
an  elementalist  nor  an  organismalist,  but  an  eminent  sub- 
jective idealist.  Josiah  Royce  is  the  student  who  has  per- 
formed this  service.  Stated  in  a  single  sentence,  the  ad- 
vance he  has  made  toward  discovering  the  union  consists 
in  the  recognition  of  certain  fundamental  resemblances  be- 
tween some  of  the  very  highest  operations  of  man's  mind 
and  the  pure  tropistic  operations  of  lower  animals. 

Royce's  contribution  to  this  subject  is  contained  in  his 
Outlines  of  Psychology,  and  nowhere  else  so  far  as  I  know. 
From  the  preface  of  this  book  I  gather  the  following,  partly 
by   way   of   quotation   and   partly    from    obvious   inference. 


Psychic   I  nt  eg  ration  221 

"To  my  mind,"  says  Royce,  "tin  interesting  side-light  1ms 
been  shed  u})()n  the  well-known  controversies  between  the 
associationists  on  the  one  liand,  .•uid  Hie  school  of  Wiiiidt 
and  the  partisans  of  'mental  activities'  generally,  on  the 
other,  by  the  stress  that  Professor  Loeb  has  recently  laid 
upon  the  part  that  what  he  calls  'tropisms'  play  in  the  hfe 
of  animals  of  all  grades."  ^ 

Then  after  telling  in  a  few  sentences  what  tropisms  are, 
Royce  continues:  "Now  it  is  especially  notable  that  the  'tro- 
pisms' of  Loeb  are  not,  like  the  'reflex  actions'  of  the  theories, 
modes  of  activity  primarily  determined  by  the  functions  of 
specific  nerve-centres.  Furthermore,  they  are  more  general 
and  elemental  in  their  character  than  are  any  of  the  acquired 
habits  of  an  organism."  (At  this  })oint  Royce  takes  up,  for 
a  moment,  a  matter  to  one  side  of  my  main  purpose,  namely 
the  problem  of  "self-activity"  and  "spontaneity" ;  so  I  ven- 
ture to  change  somewhat  the  order  and  emphasis  of  his  ar- 
gument.) "Now  it  has  occurred  to  me  to  maintain,  in  siil>- 
stance,  that  the  factor  in  mental  life  which  Wundt's  school 
defines  as  'Apperception'  .  .  .  may  well  be  treated,  from  the 
purely  psychological  point  of  view,  as  the  conscious  aspect 
or  accompaniment  of  a  collection  of  tendencies  of  the  type 
which  Loeb  has  called  'tropisms.'  "  ^ 

Then  we  have:  "Wundt  has  insisted  tliat  his  'Appercep- 
tion' is  no  disembodied  spiritual  entity.  I  conceive  that  Loeb 
has  indicated  to  us,  in  the  conce])t  of  the  'tropisni,'  how 
a  power  more  or  less  directive  of  the  course  of  our  asso- 
ciations, and  more  general  than  is  any  of  the  tendencies 
that  are  due,  in  us,  to  liabit,  or  to  specific  experience,  can 
find  its  embodiment  in  the  most  elemental  activities  of  our 
organism."  ^ 

What,  now,  is  the  bearing  of  this  idea  of  Royce's  on  the 
main  theme  of  this  chapter,  the  organism's  unity  as  mani- 
fested in  and  influenced  by  its  psychic  life?  As  an  initial 
step  toward  answering  this  question,  tlie  rcuhr  Is  asked   to 


222  The  Unity  of  the  Orgamism 

recur  to  the  chapter  on  tropistic  activities  and  their  an- 
atomical groundwork,  recalHng  that  it  was  the  special  aim 
of  our  discussion  to  show  the  inevitable  organismal  trend  of 
the  whole  doctrine  of  tropisms.  It  should  be  remembered 
also  that  trojjisms  are,  all  of  them,  probably,  beyond  ques- 
tion adaptive  in  fundamental  nature;  i.e.,  they  work  in  the 
interest  of  either  the  individual  as  a  whole  or  of  the  species 
to  which  the  individual  belongs.  The  circumstance  that 
under  occasional  more  or  less  artificial  conditions  the  ac- 
tivities of  an  animal  may  result  in  injury  to  it  or  even  in  its 
death,  is  not  proof  that  on  the  whole  those  activities  are  not 
advantageous.  A  horse  or  man  may  make  himself  sick  now 
and  then  by  taking  the  wrong  kind  of  food  or  too  much 
food,  but  this  does  not  prove  eating  to  be  useless  on  the 
whole,  or  non-adaptive. 

Another  important  thing  to  bear  in  mind  about  tropisms 
is  their  automaticity,  or  preferably  their  intrinsicality. 
They  are  rooted  in  and  partake  of  the  very  essence  of  the  or- 
ganism— so  much  so  that  they  manifest  themselves  inevitably 
when  the  right  external  and  internal  conditions  are  present, 
whether  the  general  ends  which  they  normally  serve  are  at- 
tained in  the  particular  instance  or  not.  The  flight  of  the 
moth  toward  the  flame,  even  at  the  sacrifice  of  its  life  some- 
times, is  a  manifestation  of  a  tendency  that  works,  on  the 
whole,  for  the  good  of  the  animal.  That  the  moth  follows 
the  impulse  even  to  death  merely  shows  how  tremendously 
deep-seated  this  type  of  reaction  is.  That  the  activity  may 
result  in  injury  or  death  in  a  special  case  is  just  because  the 
case  is  special,  i.e.,  it  is  a  departure  from  the  regular  con- 
ditions under  which  the  reaction-type  became  incorporated 
in  the  organization  of  the  creature.  Being  always  poised 
for  a  particular  kind  of  action,  and  having  a  supply  of  en- 
ergy to  execute  the  action,  are  unquestionably  among  the 
most  distinctive  attributes  of  animal  organisms.  Such  or- 
ganisms  are  distinguished   from  plant   organisms   not   only 


Psychic  Integration  223 

bj  the  present  fact  of  inlKrunt  activity  of  tlie  animal,  hut 
by  their  inherent  preparedness  for  acting  to  meet  new  and 
more  or  less  unusual  situations.  This  action  and  action- 
readiness  are  the  real  meaning  of  the  neuro-muscular  syst(  ni. 
All  biotic  organization  is  anticipatory  in  various  ways,  hut 
ainmals  are  almost  exclusively  anticipatory  in  action. 

It  IS  just  these  attributes  that  lloyce  recognizes  as  com- 
mon ground  between  certain  of  the  highest  psychic  activi- 
ties of  man  and  tropistic  activities.  With  this  over})lus, 
and  in  some  cases  useless  or  even  injurious  activitv  (in- 
stanced by  the  flight  of  the  moth  toward  and  around  the 
fiame),  let  us  now  pass  to  the  uj)])er  end  of  the  gamut  of 
animal  activity  for  illustrations.  A  very  few  must  suffice. 
The  first  chosen  is  one  of  exalted  creativeness  in  art. 

From  the  vast  domain  of  art  a  more  instructive  illustra- 
tion of  over-wealth  of  self-activity  can  hardly  be  found  than 
is  afforded  by  William  Shakespeare.  A  recent  investigation 
of  his  works  undertaken  with  a  view  to  finding  what  they 
tell  about  the  "native  endowments  of  the  author"  and  prose- 
cuted with  that  love  for  accurate,  exhaustive  knowledge 
which  is  the  very  soul  of  modern  science,  leads  to  the  result 
that  of  these  endowments  "the  most  outstanding  perha])s  is 
his  exuberant  vitality."  This  characteristic  of  the  man  i."> 
exhibited  in  the  "reckless  volubility  of  almost  every  cliar- 
acter,  the  piling  up  of  fancy  upon  fancy,  of  jest  upon  jest, 
the  long  embellishment  of  humor  and  foolery  and  h()rse])lay 
for  no  other  reason  than  the  delight  thev  afford."  '  And 
incidentally,  the  strict  individualism  of  this  sort  of  thing  is 
exemplified  by  one  of  these  same  Shakes])earian  characters: 
"Come,  come,"  says  Mercutio  to  l^cinolio,  "thou  art  as  hot 
a  Jack  in  thv  mood  as  anv  in  Italv.  .  .  .  Nav,  an  thrre 
were  two  such,  we  should  have  none  shortly,  for  one  would 
kill  the  other."  "What  has  Queen  Mah  to  do  with  tiu-  ac- 
tion of  the  play  of  Rojnro  and  Juliet?  Nothing;  l)ut  Mcr- 
cutio  mentions  her,  and  before  any  one  can  stop  lu'ni  he  has 


22-i  TJie  Unity  of  the  Organism 

poured  forth  fifty  lines  of  purest  fantasy.  .  .  .  Whole 
scenes,"  this  student  declares,  "exist  for  no  other  reason  than 
that  the  author's  brain  is  teeming  with  situations  and  humor 
and  infinite  jest."  ^ 

Now  I  hear  in  imagination  expressions  of  astonishment, 
rising  to  protest,  even  to  ridicule,  on  the  part  of  some  biol- 
ogists, and  to  horror  on  the  part  of  some  literateurs,  at  the 
idea  of  suggesting  that  there  is  anything  really  in  common 
between  the  two  groups  of  phenomena  here  placed  side  by 
side — the  activities  of  a  moth  and  of  Shakespeare !  For  the 
moment  I  do  no  more  in  reply  than  ask  the  reader  to  take 
cognizance  of  the  fact  that  the  whole  training  and  occupa- 
tion of  the  naturalist  consist  largely  in  comparing  all  sorts 
of  things,  inorganic  as  well  as  organic,  which  to  cursory 
observation  seem  unlike,  for  the  purpose  of  finding  whether 
closer  and  broader  examination  can  not  discover  resem- 
blances and  affinities  which  may  throw  light  on  the  ever- 
insistent  problem  of  origin  and  causal  relationship.  From 
that  procedure,  and  that  alone,  initially,  came  the  theory  of 
organic  evolution.  It  is  the  quintessence  of  the  organic 
method.  To  him  who  is  so  instructed  and  disciplined  that 
the  recognition  of  likeness  and  kinship  between,  for  example, 
the  prothallus  of  the  fern  and  the  flowering  plant,  or  between 
a  horse's  fore-foot  and  a  man's  hand,  will  receive  no  shock 
from  the  comparison.  The  intrinsic  justification  of  the 
comparison  will  be  deferred  until  we  have  a  few  other  illus- 
trations before  us.  Another  illustration  will  be  taken  from 
an  author,  J.  J.  Rousseau,  whose  activities  stand  about  mid- 
way between  art  proper  and  science  proper. 

"I  felt,"  Rousseau  says  in  his  Confessions,  "that  writing 
for  bread  would  soon  have  exhausted  my  genius,  etc.," 
Again :  "Nothing  vigorous  or  great  can  come  from  a  pen 
totally  venal."  And  finally:  "In  a  severe  winter,  in  the 
month  of  February,  and  in  the  situation  I  have  described,  I 
went  every  day,  morning,  and  evening,  to  pass  a  couple  of 


Psychic   J ntcy ration  225 

hours  in  an  open  alcove  wliich  was  at  the  bottom  of  the 
garden.  ...  It  was  in  this  pLace,  then,  exposed  to  freezing 
cold  that  without  being  sheltered  from  the  wind,  I  composed, 
in  the  space  of  three  weeks,  my  letter  to  D'AK-iulxrt  on 
theatres." 

If  this  sort  of  thing,  one  may  note  in  passing,  is  a  case 
of  "struggle  for  existence,"  the  existence  struggled  for  is 
on  the  highest  plane  of  j)sychic  life  and  not  on  the  j)lane 
of  mere  brute  continuance. 

The  only  other  example  will  be  one  of  activity  in  science 
proper,  i.e.,  "pure  intellect"  as  far  as  there  is  such  a  thing. 

The  case  of  some  man  devoting  the  best  of  his  life  to  the 
working  out  of  a  great  germal  idea — an  Aristotle,  a  Coper- 
nicus, a  Galileo,  a  Kant,  a  Darwin  will  serve  our  purpose 
best.  Of  these  we  choose  the  case  of  Danvin.  Consider  first 
the  youth  and  the  young  man  keenly  alive  to  the  flood  of 
sense  impressions  pouring  in  upon  him  from  external  nature, 
and  mentally — "internally" — "restless,"  as  Royce  would 
say,  from  an  undefined  though  strong  dissatisfaction  with 
the  stereotyped  school  and  university  curricula  and  modes 
of  dealing  with  subjects.  Later  comes  the  set  of  environ- 
mental influences  (chiefly  through  the  naturalist  Henley), 
quite  incidental  to  his  regular,  prescrilx'd  environment,  to 
which  he  responds  with  eagerness  and  effectiveness — an  al- 
most automatic  choosing  of  fields  of  intellectual  activity. 
Out  of  all  these  fragmentary  and  by-thc-way  experiences — 
"contents  of  consciousness" — there  is  organized  a  body  of 
natural  knowledge,  and  such  definiteness  and  j)n)mise  of  ten- 
dency as  to  justify  an  appointment  to  a  post  of  consider- 
able responsibility  and  unique  opportunity,  that  of  natural- 
ist to  H.  H.  Exploring  Ship  Beagle. 

During  the  voyage  and  from  the  new  and  strange  con- 
tacts with  nature  afforded  by  it,  there  arises  another  state 
of  "restlessness,"  this  time  concerning  the  origin  of  organic 
species,  the  "mystery  of  mysteries,"  as  Darwin  himsrlf  put 


226  The  Unity  of  the  Organism 

it.  A  matter  deserving  special  notice  is  that  the  truly  for- 
ward, the  creative  step  came  after,  and  was  conditioned 
upon,  a  period  of  dissatisfaction  with  the  prevalent  teaching 
on  the  subject.  Then  the  considerable  time  of  semi-con- 
structive observation  and  thinking  and  feeling  under  guid- 
ance of  the  general  surmise  that  species  arise  naturally  and 
not  supernaturally,  as  all  his  earlier  experiences — "contents 
of  consciousness" — had  taken  for  granted.  And  at  last  the 
final,  for  him,  great  conception,  the  hypothesis  of  the  "strug- 
gle for  existence"  and  "survival  of  the  fittest"  as  a  cause  of 
the  transformation  of  species.  The  suddenness  and  spon- 
taneousness  with  which  this  idea  emerged  into  consciousness 
should  be  specially  noticed.  Once  the  merest  suggestion  of 
it  hove  in  sight,  the  whole  hypothesis  formed  itself,  organ- 
ized itself,  rapidly  and  completely. 

The  sense  in  which  the  process  may  justly  be  called  spon- 
taneous is  important.  Although  we  well  know  that  the 
famous  hypothesis  was  suggested  by  the  reading  of  Malthus' 
work  on  population,  we  know  equally  well  that  the  most  es- 
sential features  of  the  hypothesis  were  not  contained  in  the 
teachings  of  Malthus.  There  was  something  genuinely  new 
in  the  hypothesis.  Out  of  the  former  total  of  experiences 
came  that  which  did  not  actually  exist  in  those  experiences. 
Although  the  hypothesis  was  clearly  a  product  of  something 
which  went  before,  it  was  a  synthetic  product  in  the  strictest 
sense,  in  essentially  the  sense  that  a  chemical  compound  is  a 
synthetic  product  of  its  interacting  elements,  the  sense  that 
the  most  distinctive  attributes  of  the  compound  can  not  be 
found  in  the  elements  taken  separately,  but  only  after  the 
interaction  has  actually  occurred. 

We  must  not  fail  to  consider  the  long  period  of  Darwin's 
strict  "self-activity"  in  collecting  evidence,  pro  and  con, 
bearing  on  his  hypothesis ;  and  the  activity  designed,  notice, 
to  ascertain  whether  or  not  there  is  a  process  going  on  in 
the  outer  world  of  plants  and  animals  corresponding  to  the 


Ps/jcJi'ic    I nf (•(/}•(! f}(ni  227 

l)roccss  he  had  conceived,  i.e.,  Iiad  pictured  in  his  "inner 
world"  of  consciousness.  'V\\v  ncnnineness  of  \\\r  individual, 
the  personal,  the  unicjue  eharaetei-  of  mental  hfc  and  nuiital 
creation  can  hardl^^  be  more  strikingly  illustrated  than  by 
such  cases  as  this  of  Darwin's  when  the  conce|)tion,  the  \\\- 
pothesis,  is  kept  to  one's  self  so  h)ni;-  in  oi'dci-  "to  prove'' 
whether  it  is  "true"  or  not. 

Now  I  want  to  call  particular  attention  to  the  indul)itable 
fact  that  these  illustrations  are  oidv  extreme  manifesta- 
tions of  attributes  which  are  universal  in  the  human  animal 
at  least.  There  is  no  normal  human  known  to  antliropolo^y 
which  has  not  some  measure,  no  matter  how  small,  ot"  creative 
impulse  in  art  and  in  science. 

As  a  conclusion  to  this  presentation  of  instances  I  nuist 
again  insist  upon  one  of  my  cardinal  points:  that  the  in- 
clh'idually  active  and  creative  power  of  the  human  oi'^anism 
on  its  psychical  side  is  not  a  whit  less  real,  less  objective,  less 
a  natural  phenomenon  to  the  natural  historian  than  is  the 
hidhiduaUij  creative  power  of  physical  growth  and  variation, 
and  reflex  and  tropistic  action.  Indeed,  the  thorough-going, 
consistent  zoological  naturalist,  the  substance  of  whose 
science  is  largely  animal  behavior  in  all  its  aspects,  can  not 
possibly  approve  the  effort  to  separate  com]iletely  the  two 
sorts  of  creation. 

First  Move  Toward  Shouing  the  Orgamsmal  Character  of 

the  Higher  Pay  chic  Life 

Now  for  the  further  scrutiny  of  such  i)sychical  facts  as 
those  typified  by  the  exam})les  i)reseided,  for  the  j)ur 
])ose  of  seeing  what  has  been  done  and  may  yt  t  l)«  done  to- 
ward brino-ino-  them  into  accord  with  the  organismal  con- 
ception,  the  pole  star  of  all  our  |)nvious  discussions.  'I'his 
examination  will  begin,  as  others  have  begun,  by  shownig 
how   elementalistic    attemi)ts    to    interpret    organic    j)henom- 


228 


The  Unity  of  the  Organism 


ena  soon  reveal  their  inadequacy  and  finally  break  down  as 
the  efforts  come  to  face  the  increasing  complexity  which 
progress  of  objective  research  always  finds  in  such  phenom- 
ena. 


Associationist  Psychology  a   Special  Case  of  Element alist 

Biology 

In  the  particular  psychical  realm  we  are  now  to  examine, 
elementalist  theory  has  appeared  most  prominently  as  what 
is  called  Associationism.  This  flourished  first  in  England  as 
the  school  of  English  Associationists,  David  Hartley,  near 
the  middle  of  the  eighteenth  century,  being  usually  consid- 
ered its  founder.  Psychologists  of  this  group  hold  that 
ideas,  which  for  them  appear  to  be  identical  with  sensations, 
are  the  "ultimate  elements"  of  psychic  phenomena.  "Ac- 
cording to  this  theory,  rigidly  carried  out,  all  genesis  of  new 
products  is  due  to  the  combination  of  pre-existing  ele- 
ments." ^  Even  the  passions,  according  to  Hartley  "must 
be  aggregates  of  the  ideas,  or  traces  of  the  sensible  pleas- 
ures and  pains ;  which  ideas  make  up,  by  their  number  and 
mutual  influence  upon  one  another,  for  the  faintness  and 
transitory  nature  of  each  singly  taken."  ^  The  "piling  up 
of  fancy  upon  fancy,  of  jest  upon  jest,  the  long  embel- 
lishment of  humor  and  foolery  and  horseplay"  which  Pro- 
fessor Manly  shows  characterize  many  of  the  Shakespearian 
plays,  would  be  explained,  according  to  this  kind  of  psy- 
chology, not  really  by  the  author  Shakespeare  but  by  the 
"aggregation,"  in  some  way,  within  him  of  ideas. 

And,  similarly,  the  works  which  in  popular  language  are 
said  to  be  by  a  Darwin,  a  Humboldt,  a  Copernicus,  an  Aris- 
totle, are  in  reality  not  by  but  merely  in  these  men.  The  men 
were  only  the  places  of  aggregation  of  the  elements — the 
ultimates — by  which  the  teaching  on  the  origin  of  species, 
on  the  general  character  of  the  earth,  on  the  solar  system, 


Psjjchic   Intrgrdfion  299 

on  the  (iL'oj)t'r  ineniiiiin-  of  cxtrinal  natiirc,  were  produced. 
Again  the  old  story  witli  which  we  have  iK'conie  faniihar:  not 
the  organism,  but  elements  of,  or  perhajjs  merelv  In  it,  are 
the  causal  explanation  of  whatever  occurrences  are  associ- 
ated with  the  organism.  It  is,  I  think,  safe  to  assume  that 
both  the  merits  and  the  demerits  of  associationi^l  psychol- 
ogy have  been  made  })atent  enough,  at  least  to  English- 
speaking  students,  by  the  writings  of  James  and  others. 

If  only  the  doctrine  of  "association  of  ideas''  can  be  satis- 
fied to  do  what  it  is  really  able  to  do,  and  not  insist  upon 
trying  to  do  what  it  can  not  do,  its  usefulness  is  great  and 
its  permanence  in  psychology  assured. 

As  indicated  above,  the  "huge  error,"  as  James  expresses 
it,  by  which  the  "whole  historic  doctrine  of  psychological 
association  is  tainted"  ^^  is  only  another  miscarriage  of  the 
elementalist  mode  of  reasoning,  and  so  is  subject  to  the 
general  type  of  criticism  which  the  reader  has  met  in  every 
chapter  in  this  book. 

In  order  to  divest  the  criticism  as  much  as  possible  of 
personal  flavor  I  shall  make  large  use  of  James'  language. 
"All  these  writers,"  says  James,  referring  to  Ilohbes,  Hume, 
Priestley,  Hodgson  and  the  later  English  associationists, 
"hold  more  or  less  explicitly  to  the  notion  of  atomistic  'ideas' 
which  occur.  In  Germany,  the  same  mythological  suppo- 
sition has  been  more  radically  grasped,  and  carried  out  to 
a  still  more  logical,  if  more  repulsive,  extreme,  by  Ilerbart 
and  his  followers,  who  until  recently  may  be  said  to  have 
reigned  supreme  in  their  native  country." 

Now  the  objection  to  the  doctrine  of  "atomistic  ideas" 
does  not  so  much  concern  the  conce])tion  of  ideas  as  atoms 
as  the  nature  attributed  to  these  atoms,  namely  in  assum- 
ing them  to  be  immutable,  and  sufiiciciit  in  their  isolate 
capacities  to  account  for  the  thought  and  other  products 
arising  from  their  "association."  The  following  two  (juota- 
tions  illustrate  the  form  this  criticism,  the  essence  of  which 


230  The  Unity  of  the  Organism 

is  now  very  familiar  to  us  as  biologists,  takes  when  it  ap- 
pears in  garments  of  a  psychologist's  making.  The  "huge 
error"  of  the  association  doctrine,  mentioned  above,  James 
explains,  is  "that  of  the  construction  of  our  thoughts  out 
of  the  compounding  of  themselves  together  of  immutable 
and  incessantly  recurring  'simple  ideas.'  "  ^"^ 

If  there  be  any  doubt  as  to  the  meaning  of  this  surpris- 
ingly un-James-like  wording,  there  certainly  can  not  be  as 
to  the  following:  "For  Herbart  each  idea  is  a  permanently 
existing  entity,  the  entrance  whereof  into  consciousness  is 
but  an  accidental  determination  of  its  being.  So  far  as  it 
succeeds  in  occupying  the  theatre  of  consciousness,  it  crowds 
out  another  idea  previously  there.  .  .  .  The  ingenuity  w^ith 
which  most  special  cases  of  association  are  formulated  in 
this  mechanical  language  of  struggle  and  inhibition,  is  great, 
and  surpasses  in  analytic  thoroughness  anything  that  has 
been  done  by  the  British  school.  This,  however,  is  a  doubt- 
ful merit,  in  a  case  where  the  elements  dealt  with  are  arti- 
ficial ;  and  I  must  confess  that  to  my  mind  there  is  something 
almost  hideous  in  the  glib  Herbartian  jargon  about  Vorstel- 
lungsmassen  and  their  Hemmungen  and  Hemmungssummen, 
and  sinken  and  erhehen  and  schwehen,  and  Verschmehungen 
and  C omplexionen.^^  ^^ 

The  long  and  short  of  the  "huge  error"  of  associationist 
psychology  is  that  ideas  are  no  such  independent,  immutable, 
simple  entities  as  the  doctrine  supposes ;  that  in  their  origin 
and  in  all  they  are,  and  all  they  do,  and  all  that  comes  forth 
from  their  association,  they  are  in  some  sort  and  measure 
dependent  upon — what?  Something.  Search  after  this 
something  has  been  a  large  motive  of  more  recent  psycho- 
logical inquiry. 

One  way  of  supplementing  and  rectifying  associationist 
doctrines  is  to  epitomize  the  shortcoming  of  these  doctrines 
in  the  statement  that  they  recognize  only  the  objective  side 
of   the   association   process,  whereas   the   subjective    side   is 


Pfsijcliic   Intccjrafion  231 

equally  ini])ortant.  'Hius  Pillsl)iiry :  **U  was  a  nv^Uci  of 
the  subjective  conditions  and  the  insistence  upon  the  ohjec- 
tive  side  of  the  problem  that  lias  led  the  Kn<rhsli  Associa- 
tional  School  into  disrepute.  'V\\v  explanations  that  they 
gave  were  true  as  far  as  thev  went,  hut  tlicir  incompleteness 
vitiated  the  conclusions  as  soon  as  tlicv  laid  claim  to  uni- 
versality." ^-  And  the  author  tlicn  shows,  convincingly 
enough,  how  two  sets  of  subjective  factors,  attentioFi  and 
choice,  play  a  large  and  im])ortant  role  in  determining  the 
"associative  train."  And  further,  "A  complete  explanation 
of  association  demands  that  both  sets  of  factors  [objective 
and  subjective]  be  taken  into  account;  to  omit  either  is  to 
fail  in  the  solution  of  the  problem."  ^^ 

Preliminary  Examination  of  Ohjecfirr  and  Subjective 

There  is  undoubtedly  a  real  gain  in  having  proved,  as 
Pillsbury  and  others  surely  have,  that  a  "side''  otlui-  than 
the  objective  in  association  does  exist.  It  is  highly  ad- 
vantageous, also,  to  have  learned  enough  about  this  other 
"side"  to  make  the  term  subjective  an  ap})ro})riate  name 
for  it.  But  any  one  coming  to  a  studj'  of  the  associational 
activities  of  the  organism's  psychic  life  as  we  have,  namely 
as  naturalists,  can  not  avoid,  if  true  to  his  traditions  and 
methods,  wanting  to  know  how  these  two  "sides,"  the  objec- 
tive and  the  subjective,  go  together — what  the  nature  is  of 
their  relation.  For  the  very  fact  that  they  are  two  sides 
of  one  thing  is  to  the  naturalist  prima  facie  evidence  that 
they  are  in  vital  organic  connection.  Kven  the  two  sides  of 
an  inanimate  thing,  like  the  earth  with  its  two  lu'misj>heres, 
have  a  relation  to  each  other  too  important  for  the  earth 
sciences  to  ignore  or  even  to  })ut  oft'  as  merely  "paralKl.'" 
But  when  it  comes  to  an  entity  like  a  live  animal,  the  cpiin- 
tessence  of  which  is  organization,  the  (jucstion  of  how  two 
of  its  "sides"  so  important  as  its  objective  and   its  subjec- 


232  The  Unity  of  the  Organism 

tive  are  related,  becomes  most  fundamental,  especially  when 
a  subject  like  that  of  ps^^chical  association  is  up  for  con- 
sideration. 

It  is,  then,  fundamental  to  our  enterprise  to  find  out  all 
we  can  about  the  connection  between  the  objective  and  sub- 
jective aspects  or  sides  of  the  organism.  It  would  be  folly 
to  expect  results  of  any  value  from  an  effort  of  this  sort 
without  having  first  given  attention  to  the  nature  of  each 
of  the  sides.  Now  the  objective  "side"  comes  to  much  the 
same  thing  as  the  physical  or  material  organism  as  we  are 
conceiving  the  "sides."  But  since  this  has  been  the  sub- 
ject of  our  whole  treatise  up  to  the  last  two  chapters,  and 
even  of  the  greater  part  of  these,  we  are  already  possessed 
of  enough  understanding  of  this  "side"  for  our  present 
purpose. 

As  to  the  subjective  "side"  the  case  is  different.  Into 
its  nature,  its  makeup  and  activities,  we  have  looked  very 
little — only  in  a  bird's-eye-view  fashion  thus  far  in  the 
present  chapter,  and  into  its  marginal  or  transitional  zone 
in  the  preceding  two  chapters.  We  are,  consequently, 
obliged  to  penetrate  further  into  the  subjective  realm  itself 
before  the  main  problem,  that  of  the  relation  between  the 
sides,  is  attacked. 

The  Essence  of  Wundtian  Apperception 

This  carries  us  back  to  tlie  point  at  which  we  brought 
Royce's  suggestion  of  a  relation  between  Loeb's  tropism 
theory  and  Wundt's  apperception  theory  into  the  discus- 
sion, the  return  to  this  point  being  for  the  purpose  of  using 
Wundt's  conception  to  induct  us  further  into  the  nature  of 
mental  life.  The  importance  of  examining  Wundt's  con- 
ception is  two-fold.  In  ihe  first  place,  we  want  to  know 
whether  or  not  it  is  genuinely  descriptive  of  man's  highest 
psychical  life.     If  it  is,  nothing  can  stand  in  the  way  of  its 


Psychic  Integration  233 

acceptance,  so  far,  by  tlie  anthropological  zoolorrjst.  But  in 
the  second  place  we  must  know  wlu'tiicr  or  not  it  carries,  as 
some  critics  believe  it  docs,  transcendi-ntal  or  supernatural- 
istic  implications.  If  this  charge  be  true  it  is  of  course  to 
this  extent  unsanctionablc  from  our  standpoint. 

Wundt's  most  concise  characterization  of  ajjperception 
which  I  have  found  is,  "The  process  by  which  any  content 
of  consciousness  is  brought  to  clear  comprehension  we  call 
apperception."  ^^ 

A  content  of  consciousness  is  any  definable  experience  wt* 
may  have.  All  consciousness  whatever  is  consciousness  of 
something  or  other.  This  "something  or  other,"  no  matter 
what,  nor  whether  regarded  as  a  whole  or  in  part,  is  a  con- 
tent of  consciousness,  according  to  my   understanding. 

What  is  most  distinctive  about  Wundt's  characterization 
may  be  regarded  as  centering  around  the  word  clear.  When 
a  particular  content  gets  itself  into  the  lime-light  of  con- 
sciousness— when  it  becomes  the  center  of  attention — the 
process  by  which  it  does  so  is  apperception.  On  the  other 
hand,  the  process  by  which  contents,  though  brought  into 
consciousness,  come  only  into  its  outer  zones  or  edges,  and 
do  not  monopolize  attention,  is  perception.  Though  tiiis 
getting  of  a  content  into  clearness  in  consciousness,  this 
monopolizing  of  attention,  may  take  place  passively  or 
actively  so  far  as  the  mind  as  a  whole  is  concerned,  the  ac- 
tive way  seems  to  be  the  more  distinctive  and  important,  at 
least  for  mental  life  as  a  whole. 

It  is  apparently  this  positive  activity  of  appcrce])tion, 
directed  toward  making  ])articular  contents  of  conscious- 
ness clear,  which  has  brought  criticism  upon  Wundt's  con- 
ception. "Wundt  talks,"  says  Pillsbury,  ''almost  as  if 
there  were  a  faculty  or  force  of  apperception,  something 
behind  and  superior  to  consciousness,  which  brings  about 
the  change  in  clearness  of  the  impressions.  There  is  in 
the  brain  a  definite  centre  of  ap])erception,  and  in  conscious- 


234 


The  Unity  of  the  Organism 


ness  a  force  very  closely  related  to  will,  that  in  and  of  itself 
chooses  certain  ideas  for  elevation  to  the  high  places  of 
consciousness,  and  equally  arbitrarily  rejects  others."  And 
then  follows  this  in  Pill8bur3^'s  criticism,  which  brings  out 
unmistakably  its  real  purport :  "It  is  very  much  like  the 
self-conscious  unity  of  apperception  of  Kant,  which  gives 
the  final  form  and  order  to  the  various  disconnected  elements 
of  the  mind,  and  is  in  so  far  something  inexplicable,  a  factor 
in  experience  that  must  be  assumed  without  any  further 
discussion  of  its  nature,  origin,  or  laws  of  action."  ^^  A 
suspicion,  obviously,  that  the  transcendentalism  of  Kant 
broods  over  Wundt's  theor}'.  As  to  tlie  justification  of  this 
suspicion  we  need  not  be  concerned  here.  Enough  for 
us  at  this  point  to  recognize  that  from  the  standpoint  of 
description  as  natural  history  practices  the  art,  or  aims  to 
practice  it,  Wundt's  account  of  the  way  the  mind  works  in 
a  vast  range  of  its  activities  seems  true,  and  as  far  as  it 
goes  is  satisfactory. 

Not  only  the  matter  of  clearness  of  the  contents  of  con- 
sciousness, but  their  makeup  as  well  is  important.  Al- 
though "psychical  elements"  figure  largely  in  Wundt's  sys- 
tem, one  finds  no  intimation  that  the  whole  mind  and  its 
contents  can  be  "explained"  by  reducing  them  to  "ultimate 
elements"  after  the  familiar  manner  of  elementalist  explana- 
tion. "All  the  contents  of  psychical  experience,"  Wundt 
says,  "are  of  a  composite  character."  And  it  follows  from 
this  that  ''psychical  elements,  or  the  absolutely  simple  and 
irreducible  components  of  psychical  phenomena,  are  the 
products  of  analysis  and  abstraction."  ^^ 

The  two  words,  "analysis"  and  "abstraction,"  need  par- 
ticular consideration.  The  psychical  elements  found  by 
analysis  do  not  exist,  as  such,  in  nature.  Analysis,  in  this 
case,  is  logical  or  thought-analysis,  and  not  objective  analy- 
sis. We  should  do  well  to  recall  what  was  said  in  the  dis- 
cussion of  reflexes,  namely  that  the  "simple  reflex,"  though 


Psijchic   I ntcy ration  }i.'J5 


legitimate  and  useful  as  an  aid  to  i.derprcting  the  plunnm- 
ena  of  reflex  nerve  action,  lias  no  actual  cxistcncr  in  nature. 
It,  like  the  "psychical  element,^'  i>  .ui  abstraction.  This  is 
only  another  way  of  saying  that  psychical  d.ni.nt^  are 
what  they  are  because  they  are  parts  of  the  mind  as  a  whole, 
just  as  we  have  seen  over  and  over  again  physical  elements 
of  the  body  are  what  they  are  because  they  are  i)arts  of  the 
body.  "The  specific  character  of  a  given  psychical  process 
dejjends  for  the  most  part  not  on  the  nature  of  its  elements 
so  much  as  on  cheir  union  into  a  composite  ])sychical  com- 
pound. Thus,  the  idea  of  an  extended  body  or  a  rhvthm,  are 
all  specific  forms  of  psychical  experience.  But  their  charac- 
ter as  such  is  as  little  determined  by  their  sensational  and  af- 
fective elements  as  are  the  chemical  properties  of  a  comjjound 
body  by  the  properties  of  its  chemical  elements.  Specific  char- 
acter and  elementary  nature  of  psychical  processes  are,  ac- 
cordingly, two  entirely  different  concepts."  ^'^ 

We  must  not  miss  an  essential  point  in  this,  that  since 
with  psychical  elements  just  as  with  chemical  elements  we 
never  know  exactly  what  or  how  nuich  each  ])articular  ele- 
ment contributes  to  the  compound,  we  are  obliged  to  con- 
ceive the  attributes  of  the  compounds  as  pertaining  to  the 
elements  collectively  even  before  the  compounding  lias  K-en 
done. 

So  it  comes  about  that  because  all  contents  of  conscious- 
ness are  fundamentally  com})osite,  but  are  also  resolvable  into 
components  of  various  grades  of  complexity,  synthesis  and 
analysis  have  a  prominent  place  in  tlu'  \\'un(ltian  system. 
Of  these,  synthesis  is  the  more  definitive  and  fundamental, 
since  it  enters  into  the  verv  nature  of  c()ii>(inii>ness  itself. 
Consciousness  is,  according  to  \\'undt,  the  ''inter-connec- 
tion of  psychical  compouncis.""  "It  is  the  name  for  the  gen- 
eral synthesis  of  ])sychical  processes,  in  which  synthesis  the 
single  com})ounds  are  markid  off  as  more  intimate  combina- 
tions." ^^     The  meanino-  of  this  is  made  clearer  bv  the  state- 


236  The  Unity  of  the  Organism 

ment  that  unconscious  states  like  deep  sleep,  faint,  and  so 
on,  are  the  interruption  of  these  interconnections. 


Remarks  On  Analysis  and  Synthesis 

This  brings  us  to  where  we  can  see  the  important  distinc- 
tion between  an  aggregation  and  a  synthesis — in  a  psychical 
sense  particularly — and  hkewise  between  a  fragmentation 
and  an  analysis.  No  mere  aggregation,*  as  of  ideas  or  emo- 
tions, would  make  consciousness.  Only  a  synthesis  of  con- 
stituents can  do  that.  And,  contrariwise,  while  the  mere 
severance  of  the  synthesized  components  produces  uncon- 
sciousness, an  analysis  of  them  results,  not  in  unconscious- 
ness, but  in  a  consciousness  of  the  constituent  parts  of  the 
contents  of  consciousness.  The  essence  of  consciousness  is 
unitariness — integratcdness,  in  our  general  terminology — as 
regards  the  contents  of  an  individual  organism's  psychical 
nature,  so  that  whatever  analytical  processes  the  mind  per- 
forms must  move  within  the  bounds  of  its  own  unitariness 
or  integratcdness.  Were  we  to  conceive  the  analytic  opera- 
tions of  the  mind  to  exceed  or  even  quite  to  equal  its  synthetic 
operations,  we  should  have  to  conceive  it  as  utterly  negating 
consciousness,  i.e.,  as  destroying  itself.  A  man  could  ana- 
lyze his  own  mind  in  an  elementalist  sense  only  by  suiciding. 
In  other  words,  he  could  never  do  it,  simply  because  he 
would  have  killed  himself  by  the  very  process  of  analyzing 
before  he  had  completed  his  job. 

These  remarks  on  the  distinction  between  synthesis  and 
aggregation,  and  between  analysis  and  fragmentation,  are 
not  quite  what  Wundt  sa^^s.  They  go  somewhat  beyond  his 
actual  expression,  but  are  legitimate  inferences,  I  am  quite 
sure,  from  his  discussion  as  a  whole.  And  they  help  us  toward 
what  we  want  to  accomplish,  namely  to  discover  still  more 

*  Recall  our  previous  remarks  on  this  subject,  e.g.,  pp.  183  and  268, 
and  also  the  quotation  from  Hartley,  p.  228. 


Psijchic   Integration  237 

than  Rojcc  discovered  about  the  relation  between  ai)percep- 
tive  processes,  as  Wundt  conceives  them,  and  the  processes 
known  as  tropisnis. 

Anticipating  our  results,  and  stating  them  in  the  most 
general  terms  possible,  we  may  say  that  the  "apperceptive 
synthesis"  of  Wundt,  and  what  may  be  called  troj)istic  syn- 
thesis, have  a  common  ground  in  the  kind  of  synthesis  which 
is  the  very  essence  of  that  kind  of  organization  to  which  the 
term  life  is  ap})lied.  To  be  alive  is  to  be  an  organic  in- 
dividual; to  be  an  organic  individual  is  to  be  an  indi- 
vidual that  perpetually  synthesizes  itself  from  substances 
extraneous  to  itself  (food,  in  the  narrower  sense,  and 
oxygen)  ;  and  to  be  a  psychically  endowed  individual 
is  to  be  an  individual  which  in  addition  to  synthesizing 
a  physical  nature  from  the  substances  mentioned,  synthesizes 
a  psychical  nature  from  physical  and  chemical  contacts  and 
interactions  between  the  individual  and  the  external  world, 
the  physical  contacts  being  called  stimuli.  ^'iewing  the 
matter  thus,  it  is  seen  to  be  highly  probable  that  in  its  ulti- 
mate essences  the  dependence  of  the  psychical  nature  of  the 
organism  on  stimuli  is  connected,  directly  and  inseparably, 
wdth  the  dependence  of  its  material  nature  on  material  nu- 
triment. 

We  should,  I  think,  be  surprised  were  a  demonstration 
to  be  produced  that  psychic  life  has  as  little  connection  with 
metabolic  processes  as  the  text-books  of  psychology  would 
lead  one  to  suppose.  Every  modern  psychologist,  like  every 
modern  biologist,  accepts,  unquestioningly  I  presume,  the 
conception  that  in  some  icay  the  psychic  life  is  no  less  de- 
pendent on  the  nutritive  substances  and  processes  than  is 
the  physical  life.  Yet  that  "some  way"  ai)pears  to  Ik^  gen- 
erally regarded  as  so  remote  and  obscure  as  to  be  U^vond  the 
reach  of  profitable  treatment  by  psychological  science,  judg- 
ing from  the  considerable  number  of  standard  text-lM)oks 
which  I  have  consulted  on  the  point.     In  only  one  of  these 


238 


The  Unity  of  the  Organism 


{Elements  of  Physiological  Psychology,  by  Ladd  and  Wood- 
ward) do  I  find  the  word  "metabolism"  in  the  index. 

Our  task  may  then  be  restated  as  that  of  making  out 
more  fully  and  clearly  than  Royce  did  the  connection  be- 
tween apperception  and  tropisms,  which  is  involved  presum- 
ably in  the  whole  problem  of  organic  synthesis  from  its 
highest  manifestations  in  psychic  synthesis  to  its  lowest 
manifestation  as  metabolic  synthesis ;  of  bringing  to  more 
specificity  the  general  statement  made  above.  But  such 
statements  are  altogether  too  sweeping  and  abstract  to  satis- 
fy scientific  description  and  explanation  in  our  day.  A 
chapter  must  now  consequently  be  devoted  to  making  them 
more  definite. 

REFERENCE  INDEX 


1.  Hobbes 

2.  Hobbes 

3.  Hobbes 

4.  Royce 

5.  Royce 

6.  Royce 

7.  Manly 

8.  Manly 


15 
21 

25 
ix 

X 

xi 
3 
4 


9.  Baldwin    I,  80 


10.  James    ('90)    1,553 

11.  James    ('90)    I,  603 

12.  Pillsbury     106 

13.  Pillsbury   112 

14.  Wundt   229 

15.  Pillsbury     270 

16.  Wundt  *. 32 

17.  Wundt   33 

18.  W^undt     223 


Chapter  XXIII 

ORGANIC     CONNECTION      HirrWKEN      PIIVSK  AL 

AND   PSYCHICAL 

A  Still  Closer  Look  at  the  Organistnal  Nature  of  Tropisms 

TTAVING  selected  tropisms  as  a  strategic  point  in  our 
A  A  |)rogram  of  search  for  tlie  vital  connection,  if  siidi 
exists,  between  the  physical  and  tlie  psychical,  we  must  turn 
again  to  this  subject.  Our  previous  treatment  of  tlie  tro- 
pism  theory  brought  out  the  essential  organismal  character 
of  the  type  of  activity  to  which  the  term  tropism  has  been 
applied.  Tlie  result  of  that  treatment  might  Ix?  epitomized 
by  saying  that  in  so  far  as  the  theory  rests  'on  accurate 
and  adequate  description,  it  is  genuinely  organismal  and 
genuinely  sound,  but  in  so  far  as  it  rests  on  causal  ex- 
planation that  is  elementalistic  in  spirit  and  expression  it  is 
genuinely  unsound.  Our  present  aim  will  be  furthered  by 
illustrating  this  epitomized  stricture  on  the  theory  in  a  little 
different  way  from  which  we  objectified  our  criticism  in  the 
earlier  treatment. 

Every  one  familiar  witli  cunrrit  explanatory  disc'ussion 
of  tropisms  must  have  noticed  the  large  and  free  manner  in 
which  the  word  substances  is  made  use  of  In  the  explanations. 
Thus,  to  illustrate:  The  larva^  of  certain  butterflies  tnu'rge 
from  their  winter  nests  under  the  Influence  of  the  warm 
spring  sunshine,  crawl  to  the  tij)s  of  the  branches  of  some 
shrub  or  tree,  eat  the  buds  and  tender  leaves  there;  then, 
after    feeding    to    satiety    "turn    tail"    and    crawl    down    the 

239 


240  The  Unity  of  the  Organism 

branches.  This  rather  complex  and,  to  the  insects,  highly 
useful  performance  Loeb  and  others  have  proved  to  consist 
of  a  series  of  reflexes  so  interconnected  as  to  come  under 
the  tropistic  type  of  activity.  And  Loeb,  e.g.,  in  the  chap- 
ter, On  the  Theory  of  Animal  Instincts  (Physiology  of  the 
Brain)  uses  the  case  to  good  effect  in  support  of  his 
contention  that  the  traditional  instinct-and-nerve-center  ex- 
planation of  such  phenomena  is  utterly  inadequate. 

So  far,  good.  As  to  the  straightforward  presentation  of 
facts,  Loeb's  position  seems  unassailable.  But  what  about 
the  causal  explanation  of  the  facts.'^  What,  exactly,  is  it 
that  sends  the  larvae  up  the  branches.'^  What  causes  the 
eating  activities.^  What  makes  the  creatures  then  turn 
about  and  finally  sends  them  down  the  branches.^  That 
several  environmental  factors,  the  warm  weather,  the 
sunlight,  the  character  of  the  plant  buds  and  leaves,  and  so 
on,  are  involved  is  brought  out  clearly  enough.  But  what 
about  the  factors  pertaining  to  tlie  larvae  themselves.'^  The 
body-shape,  the  skin,  the  sense  organs,  the  muscles  are,  as 
was  emphasii^ed  in  the  previous  discussion  of  tropisms,  freely 
recognized  after  a  fashion  by  the  tropism  theory.  But 
deeper  still  than  these — what.^^  Chemical  substances  "ac- 
cording to  requirement,"  in  the  language  of  the  cook  books. 
Until  the  caterpillars  have  taken  food  they  are  positively 
"heliotropic,"  that  is,  literally,  are  induced  by  sunlight  to 
move  toward  the  sun,  after  the  higher  spring  temperature 
has  caused  chemical  changes  in  their  bodies  essential  to 
such  movement.  But  by  eating  to  satiety  the  chemical 
changes  essential  to  the  positive  heliotropism  are  inhibited 
and  a  negatively  heliotropic  state  comes  on.  "We  can  im- 
agine," writes  Loeb,  "that  the  taking  up  of  food  leads  to 
the  destruction  of  the  substances  in  the  skin  of  the  animal 
which  are  sensitive  to  light,  upon  which  substances  the  helio- 
tropism depends,  or  that  through  the  consumption  of  food 
the  action  of  these  substances  is  indirectly  prevented."  ^     In 


Organic  Connection  Between  Physical  and  Psychical     241 

the  total  scheme,  then,  actual  and   iniacrincd,   various  "sub- 
stances" are  indispensable. 

That  the  imaginary  constituents  constitute  a  very  im- 
portant part  of  the  exphmation  is  obvious.  This  fact  is, 
however,  not  specially  objectionable.  It  is  not  if  its  tiiu' 
character  is  never  forfrotten.  But  here  comes  the  point  1 
wish  to  make  focal  just  now.  If  imagination  is  to  be  given 
a  place  at  all  in  the  argument  it  must  have  a  larger  place 
than  Loeb  has  accorded  it.  Othei-wise  the  teachings  of 
evolution,  i.e.,  the  genetic  continuity  in  biology,  are  tacitlv 
repudiated.  Attention  has  previously  been  called,  especiallv 
in  the  chapter  on  the  organism  and  its  chemistry,  to  the  deep 
current  of  virtual  anti-geneticism  which  runs  through  physi- 
ology generally,  and  particularly  through  bio-chemistrv. 
Undoubtedly  we  can  imagine  "substances"  produced  and  de- 
stroyed in  such  a  complex  of  activities  as  that  described,  to 
meet  exactly  the  needs  of  the  larva ;  but  can  we  legitimately 
imagine  them  to  be  so  produced  and  so  destroyed  by  any 
other  means  than  just  by  the  particular  animals  in  question, 
that  is,  by  the  organisms?  Various  of  our  discussions,  but 
particularly  those  in  which  the  specificity  of  protoplasm 
have  been  dwelt  upon,  constitute  a  decisive  negative  answer 
to  this  question.  Xo  causal  ex])lanation  of  the  requisite 
"substances"  imagined  can  stop  short  of  the  organism^  alive 
and  normal,  as  an  essential  and  "causal  factor"  in  the  phe- 
nomena presented.  Causal  explanation  of  tropisms  which 
aims  to  reach  a  physico-chemical  basis  is  really  organismal 
as  well  as  are  tropisms  seen  through  the  medium  of  ]Mire 
description. 

The  Automatic  and  Anticipatory  Character  of  TrojJisni.s  una 

Other  Reflexes 

Nor  should  the  reader  fail  to  note  the  intrinsicality,  the 
adaptiveness,  and  the  anticipatoriness,  of  tropisms  as  illus- 


242  TJie  UniUj  of  the  Organism 

trated  in  this  example.  Conscious  will  and  choice  seemingly 
do  not  come  into  the  operation  at  all.  Given  the  right  con- 
ditions, internal  and  external,  the  caterpillar  goes  through 
the  concatenation  of  operations  necessary  for  its  existence, 
willy-nilly.  Moreover,  the  actions  initiated  by  the  warm 
weather,  the  larvae  being  yet  down  at  the  base  of  the  shrubs 
or  branches,  have  in  organic  prospect,  as  one  might  say,  a 
supply  of  food  peculiar  to  the  species.  And  this  supply, 
be  it  remarked,  is  several  inches  at  least,  and  several  min- 
utes at  least,  away  from  the  larva  at  the  beginning  of  its 
round  of  activities.  Its  future,  even  more  obviously  than 
its  present,  existence  is  involved  in  the  acts.  Anticipatori- 
ness  is  perhaps  the  most  conspicuous  attribute  of  the  adap- 
tiveness  of  such  activities.  C.  Lloyd  Morgan  has  well  ex- 
pressed the  truth  that  one  of  the  most  important  lessons 
to  be  learned  from  the  study  of  animal  behavior  "in  its  or- 
ganic aspect"  is  the  fact  that  "living  cells  may  react  to 
stimuli  in  a  manner  which  we  perceive  to  be  subservient  to  a 
biological  end,  and  yet  react  without  conscious  purpose — 
that   is,   automatically."  ^ 

But  from  our  examination  of  the  cell-theory  we  conclude 
that  "living  cells"  in  this  statement  ought  to  read  "living 
organisms." 

So  much  by  way  of  further  preparation,  in  the  reflex  and 
tropistic  phases  of  animal  life,  for  our  search  after  a  vital 
connection  between  the  physical  and  the  psychical.  It  will 
now  be  advantageous  to  return  to  that  supremely  important 
aspect  of  human  psychic  life  already  examined  somewhat, 
namely  that  of  Wundtian  apperception. 

A  Still  Closer  Look  at   the  Likeness  Between  Higher  Ra- 
tional Life  and  Tropisms 

As  Royce's  statement  of  the  objections  to  the  concep- 
tion of  Wundt  contains  several  points  that  will  be  useful  to 


Organic  Cormection  Betzveen  Physical  and  Psychical     243 

us,  we  reproduce  more  of  his  sentences.  ''It  has  been  ob- 
jected to  the  partisans  of  Wundt  that  the  tcriii  'appercep- 
tion,' as  thus  used,  seems  to  signify  a  factor  in  mental  life 
which  can  be  explained  neither  in  terms  of  what  we  have 
called  sensitiveness,  nor  in  terms  of  the  law  of  habit.  It 
has  also  been  objected  that  the  conception  of  a  conscious 
process,  engaged  in  influencing  its  own  states,  is  a  concep- 
tion which  confuses  together  metaph3^sical  and  psycliolog- 
ical  motives.  The  psychologist,  engaged  as  he  is,  not  in 
studying  how  Reason  forms  the  world,  but  in  observing  and 
reducing  to  rule  the  mere  phenomena  of  human  mental  life  as 
they  occur,  is  not  interested,  it  has  been  asserted,  in  a  power 
whose  influence  upon  mental  phenomena  seems  to  be  of  so 
ambiguous  a  character  as  is  that  which  the  Wundtian  'ap- 
perception' possesses."  ^ 

Again :  "This  is  the  place,"  Royce  writes,  "neither  to 
expound  nor  estimate  Wundt's  theory.  But  it  does  here 
concern  us  to  point  out  that  what  occurs  in  mind  whenever 
we  are  actively  attentive  is  attended  with  a  feeling  of  rest- 
lessness, which  makes  its  dissatisfied  mth  all  those  associa- 
tive processes  that  do  not  tend  to  further  our  current  in- 
tellectual interests.  On  the  other  hand,  the  cerebral  proc- 
esses that  accompany  active  attention  are  certainly  such  as 
tend  to  inhibit  many  associative  processes  thut  would,  if 
free,  hinder  our  current  intellectual  interests .'*  Mean- 
while, "owr  active  attention  itself  is  always  the  expression 
of  interests  which  possess  the  sam-e  elemental  character  that 
we  have  all  along  been  illustrating  in  the  foregoing  para- 
graphs. The  attentive  inventor  is  eager  about  the  beau- 
tiful things  that  he  thinks  of  while  he  is  trying  to  invent. 
The  attentive  hostess  is  eager  about  social  success.  Thr 
attentive  caged  animal  is  eager  about  whatevi*r  suggest^  a 
way    of  escape."  ^ 

The  discussion  from  which  these  sentences  are  taken  is 
contained   in   a   chapter  near   the  end   of   the  book,  entitled 


244  The  Unity  of  the  Organism 

The  Conditions  of  Mental  Initiative^  and  in  order  that  the 
reader  may  get  the  full  force  of  what  Rojce  is  talking 
about,  he  is  earnestly  recommended  to  read  the  entire  chap- 
ter. Only  thus  can  the  "foregoing  paragraphs"  mentioned 
be  adequately  appraised.  But  we  must  try,  in  our  own 
way,  to  get  the  essence  of  the  matter.  Royce's  presenta- 
tion is  his  way  of  insisting  upon  the  facts  of  psychical  life 
and  activity,  high  and  low,  which  have  given  rise  to  the 
Wundtian  conception  of  apperception,  these  facts  being 
the  indubitably  initiatory,  directive  and  selective  qualities 
of  mind  in  all  its  grades.  Furthermore,  Royce  dwells  on 
the  homogeneity,  as  one  may  express  it,  of  this  intrinsicality 
of  mental  life — its  initiative,  its  persistence,  and  its  selectiv- 
ity— with  the  individual  or  fluctuating  variations  which 
have  played  so  large  a  part  in  theorizing  about  organic 
evolution  and  heredity  during  the  Darwinian  era  of  biology. 
And  he  goes  back  still  further  in  good  modern  biological 
fashion,  and  connects  these  variations  with  organic  growth 
itself,  thus  calling  attention  to  the  fact  that  variations  of 
this  particular  sort  can  not  be  referred  to  environmental 
influence. 

At  this  point  we  may  stop,  as  biologists,  to  supplement 
Royce's  argument  by  pointing  out  that  variations  of  the 
sort  indicated,  are  referable  to  environmental  influence  only 
in  the  sense  that  growth  is  so  referable.  An  organism's 
securing  and  taking  in  of  its  nutritive  substances  are  un- 
doubtedly a  kind  of  response  to  contact  with  its  environ- 
ment, and  in  that  broad  sense  growth  may  be  said  to  be 
due  to  environmental  influence.  If  the  organism  had  no 
nourishment,  if  it  received  no  environmental  influence  of  this 
kind,  it  certainly  would  not  grow.  At  the  same  time,  since 
the  organism  manages  somehow  to  build  a  great  variety  of 
tissues  and  organs  out  of  one  and  the  same  supply  of  nour- 
ishment; that  is  in  response  to  one  and  the  same  "environ- 
mental  influence"    (as   we   are   agreeing  to   use   the   phrase 


Organic  Connection  Between  Physical  and  Psychical     245 

here)  there  is  no  course  open  but  to  recognize  that  the  or- 
ganism is  a  very  important,  because  indis})ensable,  factor  in 
its  own  growth  and  differentiation.  "Self  differentiation," 
so  far  as  the  whole  organism  is  concerned,  is  a  fact  than 
which  no  other  in  the  whole  domain  of  biology  is  better  es- 
tablished. Indeed,  self  differentiation  is  really  a  special 
form  of  self  growth  and  surely  no  one  would  contend  that 
environmental  influence  is  more  than  an  essential  factor  in 
the  growth  of  an  organism.  To  hold  it  to  be  a  complete 
explanation  of  the  phenomenon  would  be  too  manifestly  ab- 
surd to  receive  serious  consideration.  It  would  be  to  con- 
tend, in  effect,  that  one  of  the  processes  of  the  organism 
(its  growth)  is  more  than  all  the  processes  of  the  whole  or- 
ganism. But  since  most  if  not  all  variation  depends,  either 
directly  or  indirectly,  upon  growth,  what  is  more  natural 
than  that  the  living,  growing  organism  should  display  much 
self  variation.^ 

That  such  variations  are  among  the  most  common  phe- 
nomena presented  by  organic  beings,  there  is  no  shadow  of 
doubt  to  any  one  who  views  the  problem  broadly  and  crit- 
ically, and  with  no  domineering  preconceptions  as  to  what 
ought  to  be  and  ought  not  to  be ;  who,  in  other  words,  views 
the  organic  world  as  a  natural  historian,  guided  by  the 
mandate  "neglect  nothing,"  instead  of  as  a  physicist  in  the 
mathematico-laboratory  sense,  guided  by  the  mandate  "neg- 
lect everything  which  can  not  be  made  to  conform  to  gen- 
eral mathematically  statable  law." 

These  remarks  about  the  relation  of  mental  activity  to 
growth,  differentiation  and  variation  of  the  organism,  and 
to  environmental  Influence  would  apply  throughout,  muta- 
tis mutandis,  to  troplstic  activity. 


246 


The  Unity  of  the  Organism 


A  Still  Closer  Description  of  the  Suhrational  Moiety  of 

Psychic  Life 

And  this  brings  us  to  where  our  final  return  may  be  made 
for  purely  descriptive  and  comparative  purposes,  to  the 
subrational  moiety  of  psychic  life,  the  purpose  of  the  re- 
turn tliis  time  being  to  characterize  this  moiety  as  faith- 
fully but  as  briefly  as  possible  on  the  basis  of  the  total  re- 
sults of  researches  in  the  field  up  to  the  present  time.  So 
bulky  and  varied  are  these  results  that  to  examine  them 
exhaustively  is  hardly  possible  for  any  one  person  even 
though  he  be  a  specialist  in  the  field.  Much  less  possible  is 
it,  then,  for  a  general  zoologist  to  make  such  an  examina- 
tion. Nevertheless  it  is,  I  believe,  possible  to  give  an 
epitome  of  the  present  state  of  knowledge  that  shall  be  true 
in  all  fundamental  respects  and  liighly  significant  for  our 
enterprise. 


Remarks  on  the  Classes  of  Suhrational  Life 

In  giving  this  epitome  we  shall  not  try  to  maintain  a  sharp 
distinction  between  reflexes,  whether  of  the  tropistic  or  any 
other  type,  and  instincts.  To  begin  with,  as  always,  when 
a  large  and  intensively  cultivated  domain  of  science  is  en- 
tered for  the  purpose  of  extracting  from  it  its  most  certain 
major  results,  we  may  take  it  for  granted  that  the  ex- 
tremists touching  any  portion  of  the  field  over  which  di- 
vergence is  wide  and  warm,  are  unsafe  guides  for  the  gen- 
eral student.  Thus,  the  student  who  enters  the  realm  of 
animal  behavior  for  such  a  purpose  as  that  for  which  we 
are  now  entering  it  soon  sees  that  those  specialists  who  find 
nothing  but  tropisms,  and  these  of  the  most  uncompro- 
mising sort,  in  the  activities  of  much  of  the  animal  king- 
dom,  are  not   the   ones   to   whose   guidance  he   can   entrust 


Organic  Connection  Between  PJif/sical  and  Psychical     247 

Iiiniself,   no   matter   liow   voluminous,   and   perhaps   excellent 
in  quality,  their  experimental  researches  may  be. 

For  instance,  such  a  view  as  that  of  Bohn's,  according  to 
which  the  word  instinct  ought  to  be  eliminated  from  the 
terminology  of  science  ''as  a  legacy  of  the  past,  the  middle 
ages,  the  theologians  and  the  metaphysicians,"  ^  is  so  obvi- 
ously unjustifiable  to  any  well-informed  zoologist  as  to 
make  him  suspicious  of  such  a  writer  all  along  the  line, 
especially  wherever  his  judgment  and  scientific  poise  are  im- 
plicated. 

This  question  of  the  reality  of  instincts  I  use  to  Illus- 
trate the  peril  to  the  general  student  of  the  unpoised  spe- 
cialist, because  it  is  germane  to  the  present  discussion.  In 
general  zoology  the  type  of  animal  behavior  to  which  the 
term  instinctive  is  applied  is  not  less  conspicuous  and  real, 
to  say  the  least,  than  is  the  type  described  as  tropistic. 
For  an  experimentalist  to  come  out  of  his  laboratory  and 
tell  a  broadly  experienced  entomologist  or  ornithologist,  for 
example,  that  the  familiar  achievements  of  young  insects 
of  many  species,  and  of  numerous  young  birds  should  not 
be  called  instinctive  because  (as  the  experimenter  asserts) 
they  are  reducible  to  the  tropistic  or  ])erchance  the  simple 
reflex  type  of  reaction,  may  justly  be  characterized  as  sci- 
entific impertinence.  It  is  as  though  an  cmbryologist,  hav- 
ing discovered  that  a  bird's  wing  is  the  genetic  counterpart 
of  a  salamander's  forelimb,  should  Instruct  the  ornitholo- 
gist that  it  is  wrong  for  him  to  call  the  bird's  wing  a  wing, 
because  the  member  may  be  reduced  to  a  lower  type  of  limb. 
Unquestionably  the  experimental  specialist  often  pro- 
duces results  which  necessitate  changes  in  the  general  zo- 
ologist's conceptions  and  nomenclature.  Jkit  it  is  not  his 
province  to  take  into  his  own  hands  the  re\ision  of  the 
fundamental  terms  of  zoology.  Any  one  moderately  In- 
structed in  the  history  of  zoology  knows  that  ''instinct''  is 
a  hardly  less  well-grounded  zoological  term  than  "birth"  or 


248  The  Unity  of  the  Organism 

"intelligence,"  or  many  another  indispensable  term. 

Our  inquiry  is  not  as  to  whether  there  are  such  things  as 
instincts,  but  how  they  operate  and  what  they  signify  for 
the  animals  possessing  them.  Perhaps  of  highest  interest 
to  us  is  the  fact  that  innumerable  instincts,  if  indeed  not  all, 
are  as  indubitably  hereditary  as  are  any  animal  endowments 
whatever.  This  comes  out  especially  convincingly  in  those 
numberless  cases  where  the  instinctive  operations  develop 
strictly  pari  passu  with  the  anatomical  development  of  the 
young,  there  being  absolutely  no  opportunity  for  them  to 
learn,  even  subconsciously. 

Take  as  an  example  the  crustacean  Amphithoe  longimana, 
in  which  Holmes  compared  in  detail  the  activities  of  the 
newly  hatched  young  with  those  of  the  adult.  "Amphithoe 
lives  in  tubular  nests  which  are  usually  lodged  among  sea 
weed.  The  nests  are  somewhat  longer  than  the  animal, 
and  are  spun  of  a  web-like  material  into  which  bits  of  sea 
weed  are  often  incorporated  which  help  to  conceal  the  oc- 
cupant. In  its  nest  Amphithoe  lies  in  wait  for  prej,  ready 
to  dart  out  upon  any  small  creature  which  touches  the  ends 
of  its  long  antennae. 

"The  activities  of  the  adult  Amphithoe,  with  the  excep- 
tion of  those  concerned  in  reproduction,  are  almost  ex- 
actly paralleled  by  those  of  the  young.  I  have  taken  the 
eggs  from  the  maternal  brood  pouch  shortly  before  hatch- 
ing and  kept  them  isolated  in  individual  dishes.  For  some 
time  after  emerging  from  the  egg  the  young  were  weak  and 
had  imperfect  control  of  their  movements,  which  were  jerky 
and  irregular.  Soon  the  minute  creatures  would  crawl  and 
swim  much  like  the  adults,  and  the  next  day  they  began 
constructing  nests  which  were  the  same  shape  as  those  formed 
by  their  parents."  Then  comes  a  part  of  the  description 
to  which  the  reader's  special  attention  is  called  because  it 
brings  out,  partly  by  implication,  a  richness  of  detail  in  be- 
havior which  defies  full  expression,  and  which  every  care- 


Organic  Connection  Between  Physical  and  Psychical     249 

fully  observing  zoologist  knows  to  be  characteristic  of  the 
activities  of  nearly  all  animals.  Especial  attention  is  in- 
vited to  this,  because  this  elusive  wealth  of  behavior  is 
usually  overlooked  by  the  cursory  observer  on  the  one  hand 
and  by  the  experimentalist  on  the  other.  "The  attitudes  in 
the  nest,"  Holmes  writes,  "the  waving  of  the  antenna?,  the 
beating  of  the  swimmerets,  the  restless  movements  of  the 
legs  and  mouth-parts,  springing  after  food,  belligerency 
toward  passers  by,  the  little  unobtrusive  signs  of  timidity, 
the  reversal  of  position  in  the  nest  on  the  approach  of 
danger  and  the  general  behavior  outside  of  the  nest,  were, 
on  the  next  day  after  hatching,  almost  exactly  the  same  as 
in  older  individuals.  The  only  differences  in  behavior  were 
due  to  the  feebleness  of  the  young  and  their  imperfect  con- 
trol of  their  movements."  One  never  reads  a  description 
like  this  by  a  typical  experimentalist,  especially  if  he  be  a 
pure  tropist,  or  by  a  meagerly  trained  zoologist !  Then  the 
final  statement :  "The  young  are  hatched  with  all  the  in- 
stincts necessary  fully  to  equip  them  for  the  business  of  life. 
No  experience  is  necessary  to  teach  them  what  is  advan- 
tageous for  them  to  do."  ^ 

The  impossibility  should  be  noticed  of  drawing  a  sharp 
line  in  this  description  between  instinctive  and  purely  reflex 
acts.  "Reversal  of  position  in  the  nest  on  approach  of 
danger"  is  clearly  instinctive.  But  "beating  of  swimmer- 
ets," and  especially  the  "restless  movements  of  the  legs" — 
are  these  instinctive  or  wholly  reflex?  Probably  they  arc 
reflex,  though  the  leg  movements  may  well  be  partly  in- 
stinctive. A  whole  volume  of  examples  as  unquestionable 
as  this  could  be  compiled,  and  all  groups  of  animals  from 
mammals  down  to  worms  at  least  would  be  represented. 


250  The  Unity  of  the  Organism 

Four  Certainties  About  the  Adaptiveness  of  Subrational 

Psychic  Activities 

Concerning  the  purposefulness  or  adaptiveness  of  activi- 
ties of  this  general  type,  I  think  four  things  may  be  re- 
garded as  absohitely  certain. 

Generally  Us,eful  to  Individual  and  to  Species 

First,  a  vast  majority  of  them  are  recognizably  contribu- 
tory to  the  perpetuity  of  both  the  individual  in  its  normal 
life,  and  of  the  species.  But  for  them  neither  individual 
nor  species  would  continue  to  exist.  This  is  so  obvious  that 
further  remark  upon  it  is  unnecessary. 

Many  Useful  to  Species  Primarily 

Second,  in  a  large  number  of  instances  particular  acts 
by  particular  indi\4duals  are  in  the  interest  of  the  species 
primarily  and  of  the  individuals  only  secondarily  or  not  at 
all.  This  is  shown  most  conclusively  in  cases  like  that  of 
several  species  of  salmon,  where  the  individual  normally  goes 
through  activities  which  secure  the  continuance  of  the  spe- 
cies but  which  end  in  the  death  of  the  individual.  A  large 
and  varied  number  of  cases  of  this  type  occur,  especially 
among  insects.  But  the  supremacy  of  species  over  individ- 
ual needs  appears  under  various  other  forms.  Thus  almost 
certainly  such  tropistic  activities  as  that  of  the  moth  going 
to  its  death  or  injury  in  the  flame  is  of  this  sort.  This 
case  may  be  stated  in  general  terms  thus :  Owing  to  lack 
of  any  ability  on  the  part  of  an  individual  to  modify  its  in- 
herited mode  of  action  to  meet  a  special  situation,  it  acts  in 
the  old  way  even  though  the  new  situation,  while  in  gen- 
eral like  the  old,  yet  differs  from  it  enough  to  make  it  peril- 
ous to  the  individual  if  it  acts  unmodifiedly  in  the  old  racial 


Organic  Connection  Between  Physical  and  Psychical     ^51 

way.  The  preeminently  racial  utility  and  hereditary  char- 
acter of  instincts  is  certainly  one  of  the  most  interestinrr 
things  about  them  for  the  present  discussion. 


Variability  of  Suhralional  Activities 

The  third  certainty  about  reflex  and  instinctive  activities 
is  that  they  are  b}-  no  means  so  stereotyped  and  invariable 
as  older  cursory  observation  or  as  much  theorizing,  espe- 
cially about  tropisms  and  instincts,  has  held  them  to  be. 
Darwin,  in  the  notable  chapter  on  Instinct  in  The  Origin  of 
Species,  was  the  first  to  attack  seriously  the  notion  of  such 
invariability  in  dealing  with  instincts.  He  undertook  to 
show  that  the  instinctive  type  of  activity  is  subject  to  vari- 
ation just  as  are  all  other  aspects  of  animal  life. 

A  telling  set  of  recent  investigations  under  this  head  is 
by  the  Peckliaiiis.  That  on  tlie  solitary  wasp,  AnimopliiUi 
ui'nariay  is  particularly  to  the  point  because  the  earlier 
writers  had  used  its  habits  of  paralyzing  caterpillars  by 
stinging  them  and  storing  them  up  as  food  for  its  young  to 
illustrate  the  undeviating  and  unerring  character  of  in- 
stincts. But  the  extensive  studies  of  these  entomologists 
led  them  to  write :  "The  one  preeminent,  unmistakable  and 
ever-present  fact  is  variability.  Variability  in  every  par- 
ticular— in  shape  of  the  nest  and  the  manner  of  digging 
it,  in  the  condition  of  the  nest  (whether  closed  or  open) 
when  left  temporarily,  in  the  method  of  stinging  the  prey, 
in  the  degree  of  malaxation,  in  the  manner  of  carrying  the 
victim,  in  the  way  of  closing  the  nest,  and  last,  and  most  ini- 
])ortant  of  all,  in  the  condition  produced  in  tlie  victims  l)v 
stniguig.     ' 

No  present-day  authority  so  far  as  I  know  contends  that 
instincts  operate  in  a  hard-and-fast  manner  comparable  to 
the  workings  of  any  man-made  machine.  They  are  now 
universally   recognized    to   be    subject   to    the   same   general 


252 


The  Unity  of  the  Organism 


principles  of  variation  to  which  all  organic  phenomena  are 
subject.  Furthermore,  under  the  searching  investigation 
and  criticism  of  numerous  workers,  notably  H.  S.  Jennings 
and  his  followers,  the  tropism  theory  has  been  deprived,  for 
most  biologists,  of  its  inorganically  mechanistic  character. 
The  principles  of  "random  movements,"  "avoiding  reac- 
tions," "trial  and  error,"  and  others,  are  thoroughly  estab- 
lished and  the  recognition  of  them  may  be  said  to  have  so 
modified  the  doctrine  of  tropisms  as  to  make  it  one  of  or- 
ganic mechanism  rather  than  of  inorganic  mechanism — as  it 
virtually  would  be  according  to  the  thoroughgoing  elemen- 
talistic  conception  of  it.  The  "mechanistic  conception  of 
life,"  one  may  remark,  has  very  much  to  commend  it  if  only 
the  machines  conceived  are  recognized  to  be  alive.  My  re- 
marks under  this  head  *  may  be  consulted  by  the  reader 
who  wishes  to  follow  this  point. 

What  is  meant  by  random  movements  is  made  clear  by 
the  following:  "In  the  earthworm  and  the  larvae  of  blow- 
flies which  are  negatively  phototactic  it  has  been  shown  by 
the  writer  that  movements  which  bring  the  animal  toward 
the  light  are  checked  or  reversed  and  only  those  which  hap- 
pen to  direct  the  animal  away  from  the  light  are  followed 
up.  Whatever  immediate  orienting  tendency  the  light  may 
have  in  these  cases  is  relatively  unimportant  as  compared 
with  the  element  of  selection  of  favorable  movements  in  di- 
recting the  animal  away  from  the  light."  * 

Here  it  will  be  noticed  that  the  end,  beneficial  to  the  ani- 
mal, is  reached  through  a  combination  of  orienting  reac- 
tions of  the  rigidly  tropistic  type,  i.e.,  the  type  dependent 
on  the  movement  of  the  animal  directly  toward  or  away 
from  the  source  of  light  by  the  symmetrical  plan  of  the 
body,  and  a  sort  of  reaction  in  which  the  particular  body- 
form  and  the  direction  of  light  rays  are  of  only  secondary 

*  See  "Machines,  living,"  in  the  index  of  The  Probable  Infinity  of  Na- 
ture and  Life. 


Organic  Connection  Between  Physical  and  Psychical     253 

significance.  But  tlii.s  latter  type  of  activity,  wholly  di- 
vorced from  a  direct-orienting  reaction,  and  even  from  a 
bilateral  body  s^nnmetry,  is  of  wide  application  among  the 
lower  animals.  It  was  first  brought  clearly  to  the  atten- 
tion of  biologists  by  Jennings  in  his  now  well-known  inves- 
tigations on  Paramecium  and  other  protozoans.  These  in- 
vestigations formed  the  bases  of  the  "avoiding  reaction"  and 
the  "trial  and  error"  conceptions  now  generally  recognized 
to  be  of  much  importance  in  the  behavior  of  all  animals,  es- 
pecially of  those  in  which  a  high  measure  of  bodily  activity 
occurs  but  in  which  there  is  little  or  no  intelligence.  Jen- 
nings' lucid  account  of*  his  results  in  the  chapter  Be- 
havior of  the  Infusoria;  Paramecium  (Behavior  of  Lower 
Organisms)  is  strongly  commended  to  the  reader. 

The  following  paragraph  must  suffice  for  our  reference 
to  this  w^ork.  After  describing  the  behavior  of  Parame- 
cium, Jennings  writes :  "This  method  of  behaving  is  per- 
haps as  effective  a  plan  for  meeting  all  sorts  of  conditions  as 
could  be  devised  for  so  simple  a  creature.  On  getting  into 
difficulties  the  animal  retraces  its  course  for  a  distance,  then 
tries  going  ahead  in  various  directions,  till  it  finds  one  in 
which  there  is  no  further  obstacle  to  its  progress.  In  this 
direction  it  continues.  Through  systematically  testing  the 
surroundings,  by  swinging  the  anterior  end  in  a  circle,  and 
through  performing  the  entire  reaction  repeatedly,  the  in- 
fusorian  is  bound  in  time  to  find  any  existing  egress  from 
the  difficulties,  even  though  it  be  but  a  narrow  and  tortuous 
passageway."  ®  And  this  complex  and  highly  useful  be- 
havior is  performed  by  an  organism  which,  so  far  as  the 
best  anatomical  researches  have  been  able  to  determine,  is 
entirely  devoid  of  a  nervous  s^^stem,  and  consists  of  a  single 
cell! 

But  the  "trial  and  error"  scheme  here  exemplified  is  by 
no  means  confined  to  unicellular,  non-nervous  animals,  nor 
to   experimentally  produced   conditions.      That   it   is   opera- 


254  The  Unity  of  the  Organism 

tive  in  nature,  and  among  animals  with  rather  highly  de- 
veloped nervous  systems  I  shall  illustrate  by  describing 
briefly  a  performance  witnessed  by  me  some  years  ago. 
This  was  the  capture  and  engulfment  of  food  by  a  nemertean 
worm.* 

These  marine  worms  are  of  considerable  size,  some  reach- 
ing a  length  of  many  inches,  even  a  few  feet,  and  ranging 
in  thickness  from  less  than  an  eighth  of  an  inch  to  nearly  an 
inch.  Externally  they  give  the  impression  of  being  very 
lowly  in  organization,  the  body  being  devoid  of  limbs  or 
other  appendages,  and  without  segmentation.  However, 
when  they  are  examined  internally  a  surprisingly  high  grade 
of  organization  is  found,  the  muscular,  digestive,  blood  and 
nervous  systems  being  on  a  par,  probably,  with  those  of 
any  invertebrates  below  the  crustaceans  and  insects.  The 
nervous  system,  particularly  the  brain,  is  relatively  large, 
though  not  differentiated  into  diverse  ganglionic  masses  and 
connecting  strands  to  the  extent  found  in  jointed  worms. 
The  creatures  are  poorly  equipped  with  external  sense  or- 
gans, there  being  no  tentacles  nor  any  certain  olfactory  or 
auditory  organs.  And  eyes,  when  present,  are  so  minute 
and  simple  as  to  be  without  power  of  sight  in  the  ordinary 
sense;  almost  certainly  they  are  mere  light-perceiving  or- 
gans. 

The  most  distinctive  anatomical  feature  of  the  nemerte- 
ans  is  a  very  long  and  thin  though  muscular  and  flexible  hol- 
low tube  situated  at  the  anterior  end  of  the  animal,  which 
is  usually  carried  stowed  away  in  a  pouch  within  the  body. 
While  thus  retracted  the  tube  has  some  such  relation  to 
the  rest  of  the  animal  that  a  glove-finger  would  have  to  the 

*  Greatly  to  my  regret  I  am  unable  to  say  what  the  species  or  even  the 
genus  was  of  either  the  nemertean  or  the  annelid  here  referred  to.  The 
observation  was  made  at  the  Shumagin  Islands,  Alaska,  and  under  cir- 
cumstances that  rendered  it  quite  impossible  to  "look  up"  the  species. 
And  my  knowledge  of  the  taxonomy  of  these  groups  of  worms  is  alto- 
gether too  meager  to  enable  me  to  identify  genera  even,  offhand. 


Organic  Connection  Between  Physical  and  Fsychical     '■Zi>:j 

glove  were  it  coni2)letely  inverted  into  the  hand  of  the 
glove.  This  tube  is  used  in  the  capture  of  prey,  tlie  ani- 
mals being  carnivorous  and  highly  voracious.  The  mode  of 
employing  the  apparatus  consists  essentially  in  thrusting 
the  tube  out  with  almost  the  speed  of  lightning,  the  object 
being  to  bring  the  organ  into  contact  with  the  prey  at  many 
points.  The  lash  is  not  used  as  a  lasso  for  catching  or  as 
a  spear  for  piercing  the  prey,  but  for  paralyzing  it,  prol>- 
ably  by  a  toxic  secretion  spread  over  the  whole  surface. 
The  more  effectually  to  accomplish  this,  the  lash  is  shot  out 
at  a  victim  again  and  again. 

Now  for  the  aspect  of  the  whole  operation  of  food-taking 
which  specially  concerns  us.  Being  quite  sightless  and 
touchless  in  the  usual  sense,  the  lash  must  be  used  as  an  ex- 
ploring or  finding  as  well  as  a  paralyzing  or  killing  organ ; 
and  since  its  great  length  and  limberness  preclude  it  from 
being  used  as  an  ordinary  tentacle  is  used,  the  finding  op- 
eration is  accomplished  by  repeated  out-thrustings  of  the 
tube.  In  the  instance  witnessed  the  prey  was  an  annelid 
worm,  a  creature  well  provided  w^ith  locomotor  organs,  and 
a  crood  crawler.  On  this  account  the  victim-to-be  was  able, 
in  the  early  stages  of  the  onset,  to  move  out  of  contact  with 
the  nemertean  now  and  then.  At  such  times  the  prey  could 
be  relocated  only  by  darting  out  the  lash  at  random,  except 
as  to  general  direction.  So  it  resulted  that  many  of  the 
thrusts  missed  the  mark;  but  they  were  instantly  repeated 
with  a  little  variation  of  direction,  till  the  victim  was  lo- 
cated again.  The  whole  performance  reminded  one  of  the 
game  of  blind-man's-buff,  a  game  in  which  the  seeker  paws 
around  in  the  general  vicinity,  as  he  believes,  where  the  one 
sought  was  last  touched. 

The  effectiveness  of  the  try,  try  again  method  was  at- 
tested in  this  instance  by  the  fact  that  the  annelid  was  lilt 
and  the  paralyzing  dose  administered  times  enough  to  put 
the  annelid  into  so  helpless  a  state  that  the  nemertean  was 


^56  The  Unit!/  of  the  Organism 

finally  able  to  get  its  mouth  into  contact  with  its  prey. 
Then  the  victim,  itself  but  little  smaller  than  the  nemertean, 
disappeared  down  the  latter's  "throat"  with  almost  the 
rapidity  with  which  the  lash  was  retracted  into  and  thrust 
out  of  its  pouch.  How  much  of  this  highly  complex  per- 
formance, so  eminently  useful  to  the  nemertean,  was  purelv 
reflex,  how  much  chemotactic,  and  how  much  instinctive  r" 
And  who  will  assert  positively  that  there  was  no  trace  of 
consciousness,  even  of  intelligence,  in  it? 

An  extremely  interesting  line  of  inquiry  is  suggested  by 
cases  of  "trial  and  error"  like  this  where  at  one  extreme  the 
"errors"  are  not  much  less  numerous  than  the  successes,  and, 
at  the  other  extreme,  are  cases  in  which  the  errors  are  re- 
duced almost  to  nil.  A  type  case  of  this  last  would  be  the 
poise-and-spring  of  a  cat  after  its  prey.  With  little  doubt 
a  closely  graded  series  could  be  made  out  running  through 
from  one  extreme  to  the  other.  A  cardinal  interest  in  the 
inquiry  would  be  as  to  the  extent  to  which  the  simple  reflex, 
tropistic  reflex,  instinct,  and  intelligence  figure  in  the  dif- 
ferent grades.  Would  it  not  turn  out  that  the  gradual 
diminution  of  error  through  the  series  would  be,  generally 
speaking,  concomitant  with  the  increase  of  intelligence.?  I 
suspect  so. 

Tendency  of  Suhrational  Activities  to  Excessiven-ess 

The  fourth  and  last  certainty  about  reflex  and  instinc- 
tive activities  to  receive  attention  is  their  tendency  to  ex- 
cessiveness — their  way  of  going  beyond  what  is  necessary  or 
even  really  safe  for  the  welfare  of  the  organism.  Although 
from  several  points  of  view  this  is  one  of  the  most  impor- 
tant aspects  of  the  whole  subject,  it  has  received  surpris- 
ingly little  attention,  especially  by  the  modern  school  of  ex- 
perimental zoology. 

Probably  every  one  who  has  observed  animals  widely  and 


Organic  Connection  Between  Physical  and  Psychical     257 

thoughtfully  has  been  impressed  with  the  exuberance  of  their 
performances.  That  they  are  ever  wont  to  overdo  thlnirs, 
even  operations  which  are  wlien  done  in  measured  fashion 
absolutely  essential  to  their  existence,  is  matter  of  common 
knowledge.  Holmes  has  some  comments  under  this  head 
which  may  fitly  introduce  our  presentation.  "With  all  their 
wonderful  adaptiveness  instincts  are  far  from  ideallv  per- 
fect. Much  of  Mark  Twain's  remarks  on  the  futility  and 
imbecility,  the  wasted  effort  and  labor  at  cross  purj)oses 
shown  in  the  behavior  of  ants  may  easily  be  verified  by  anv 
observer."  ^^ 

A  common  form  taken  by  excessiveness  of  action  is  repe- 
tition. Very  many,  perhaps  all,  animals  are  notorious  re- 
peaters. A  few  out  of  the  many  available  instances  will 
suffice  to  fix  the  phenomenon  in  mind.  Some  time  ago  a 
small  whale  (probably  a  half-grown  Humpback,  Megapfera 
versabilis)  came  near  shore  at  La  Jolla,  California,  and  re- 
mained in  the  same  small  area  for  days.  While  there  it 
went  through  a  particular  set  of  movements  known  to  whal- 
ers as  "breaching"  scores  of  times,  each  set  being  exactly, 
so  far  as  one  could  see  from  the  shore,  like  every  other  set. 
The  performance  consisted,  in  this  case,  of  a  leap  out  of 
the  water,  which  carried  the  body  clear  of  the  surface  of 
the  sea,  the  direction  of  emergence  being  probably  thirty 
degrees  from  the  perpendicular.  During  the  ascent  the  ani- 
mal turned  with  a  characteristic  twist  to  the  left  and  came 
down  on  its  head  and  left  side  wdth  a  great  splash.  Once 
back  in  the  ocean  the  creature  reversed  the  course  it  was 
going  when  making  the  leap,  returned  to  some  distance  from 
where  it  had  emerged,  reversed  its  course  again,  and  re- 
peated the  leap  identically,  to  all  appearances,  even  as  to 
the  spot  of  emergence  and  direction  of  travel.  Wliy  so 
many  times  the  same  performance  in  the  same  spot?  Tliat 
is  the  problem  which  concerns  us  here.  Even  though  we 
conceive  it  to  be  somehow  adaptive — connected   in   some  in- 


258  The  Unity  of  the  Organism 

direct  way  possibly  with  feeding  or  reproduction  or  migra- 
tion or  some  other  vital  function — the  question  still  remains, 
why  so  much  of  it?  And  to  this  no  probable  or  even  ra- 
tional answer  is  forthcoming  from  the  standpoint  of  adapta- 
tion and  utility,  taking  these  terms  in  their  usual  meaning. 

Here  is  another  case  from  the  mammalia,  the  possible 
adaptive  significance  of  which  is  still  more  remote,  if  any- 
thing, than  that  of  the  behavior  of  the  whale.  Many  indi- 
vidual mice  of  the  genus  Peromyscus  being  used  by  Doctor 
Sumner  and  Mr.  Collins  in  their  researches  on  heredity  and 
environmental  influence  at  the  Scripps  Institution  take  to 
throwing  back  summersaults  in  their  cages.  The  more  com- 
mon performance  consists  in  a  run  along  the  floor  of  the 
wooden  cage  and  up  its  side  to  near  the  top,  then  a  quick, 
strong  jump  backward  clear  across  the  cage,  the  feet  being 
uppermost  during  the  first  part  of  the  leap  but  coming  to 
rights  again  by  the  time  the  landing  is  made.  Here  again 
the  question  of  why  the  mice  do  this  seemingly  useless  thing 
is  not  so  interesting  for  the  present  discussion  as  that  of 
why  they  do  it  so  much. 

The  high  flight  of  some  species  of  birds,  the  great  eleva- 
tions being  reached  by  long,  regular  upward  spirals,  would 
appear  to  come  under  the  head  of  non-adaptive,  superfluous 
action.  The  sand-hill  crane,  Grus  Tuexicana,  may  be  taken 
as  an  instance  of  a  bird  given  to  this  habit.  Surely  such 
flights  by  this  species  can  have  nothing  to  do  with  food- 
getting,  since  in  the  excursions  the  bird  is  going  directly 
away  from,  instead  of  into,  the  region  where  its  ^  food 
abounds.  It  eats  snakes,  frogs  and  other  creeping  animals, 
and  various  seeds  and  roots.  Nor  is  there  any  evidence  that 
the  flights  are  concerned  with  the  mating  function,  nor  yet 
with  migration,  though  one  might  possibly  imagine  that 
while  on  the  excursions  the  birds  learn,  after  a  fashion,  the 
topography  of  the  surrounding  regions. 

The  high-diving  and  booming  of  the  night-hawk,  Cordeiles 


Organic  Connection  Between  Physical  and  Psychical     2r)9 

viryinianus,  repeated  time  after  time  in  the  early  evening  and 
occasionally  in  midday  when  an  aijproaching  storm  cools 
the  air,  would  seem  to  be  another  j)erformance  of  the  non- 
adaptive  sort.  The  suggestion  that  this  is  a  courtship  af- 
fair can  hardly  stand,  in  view  of  the  fact  that  at  least  as 
often  as  otherwise  the  birds  which  do  it  are  entirely  alone. 
Nor  can  one  see  how  so  extensive  and  swift  a  dive,  with  so 
much  noise,  can  be  advantageous  for  the  capture  of  flying 
insects. 

And  reflect  on  the  quantity  of  movement  of  many  ani- 
mals. Can  any  one  believe  that  mammals  and  lizards  run, 
birds  and  insects  fly,  and  fishes  swim  just  exactly  so  much  as 
and  no  more  than,  they  must  in  order  to  survive.^  Would 
it  be  contended  that  the  Golden  Plover,  to  take»a  well  known 
case  of  extensive  migration,  would  certainly  succumb  in  the 
struggle  for  existence  on  anything  less  than  a  journey  from 
the  high  latitudes  of  the  northern  hemisphere  well  into  the 
southern  hemisphere  and  back,  each  year?  There  is  a  vast 
difference  between  a  necessity  for  migration  to  some  ex- 
tent and  a  necessity  for  migration  of  a  particular  quantity. 
One  of  the  great  weaknesses  of  the  natural  selection  theory 
has  been,  I  am  very  sure,  its  slight  regard  for  quantity ; 
quantity  of  need,  quantity  of  performance,  quantity  of 
benefit. 

These  examples  serve  to  illustrate  the  fact  that  among 
the  higher  animals  at  least,  much  muscular  activity  occurs 
which  is  not  at  all,  or  only  partly,  adaptive.  But  by  far  the 
more  common  occurrence  of  excessive  activity  is  in  connec- 
tion with  behavior  which  is  more  or  less  obviously  adap- 
tive. "A  o-ood  thill":  carried  to  excess,"  in  the  familiar 
phrase,  expresses  well  what  is  in  mind  here. 

This  excessiveness  of  adaptive  activity  is  naturally  more 
easily  recoo-nized  in  animals  which  are  most  easilv  observed 
and  most  active  generally.  Thus  it  Is  from  birds  and  in- 
sects that  examples  can  be  most  readily  drawn. 


260  The  Unity  of  the  Organism 

Let  the  current  view  be  accepted  that  the  song  of  pas- 
serine birds  is  associated  adaptivelj  with  the  mating  func- 
tion. Even  so,  no  one  who  has  given  careful  attention  to 
the  matter  can  have  failed  to  recognize  that  with  many 
species  much  more  singing  is  done  than  actual  pairing  and 
breeding  call  for.  I  have  kept  almost  daily  notes  for  sev- 
eral years  on  the  singing  of  the  Western  Meadow  Lark, 
Sternella  magna  neglect  a,  in  the  vicinity  of  La  Jolla.  The 
birds  are  resident  the  whole  year  through,  and  as  they  come 
familiarly  around  my  home  and  laboratory,  the  observations 
can  be  quite  full.  Although  the  breeding  time  is  restricted 
to  late  February,  March,  April,  and  sometimes  May,  there 
is  not  a  month  in  the  year  when  songs  may  not  be  heard, 
most  of  the  time  in  full  volume.  Significantly,  I  believe,  the 
song  is  at  its  ebb  during  some  weeks  just  before  the  nesting 
period  begins.  Nor  does  the  singing  of  the  males  seem  to 
be  connected  in  any  close  way  with  mating.  The  birds  do 
not  pair  off  closely  and  permanently,  even  for  the  breeding 
season.  Most  of  the  singing,  which  occurs  cliiefly  in  the 
morning  and  early  forenoon  and  again  toward  evening,  is 
done  while  the  singer  is,  more  commonly  than  otherwise, 
quite  alone  on  some  telephone  pole  or  wire.  And  the  mode 
of  singing  does  not  change  at  all  when  mating  begins.  An- 
other interesting  fact  about  the  singing  of  this  species  is 
the  considerable  range  of  temperature  and  light  conditions 
over  which  the  song  is  invariable,  so  far  as  these  factors  are 
concerned.  The  song  may  be  as  full  and  frequent  on 
cloudy,  misty  mornings  as  on  sunny  ones ;  and  over  a  con- 
siderable range  of  temperature  the  song  is  quite  independent 
of  the  particular  degree  marked  by  the  thermometer. 

While  the  song  habits  of  this  bird  are  undoubtedly  some- 
what exceptional  in  their  looseness  of  correlation  with  mat- 
ing and  with  environmental  conditions,  certain  it  is  that 
much  this  sort  of  thing  is  observable  with  several  resident 
species  which  I  have  observed.     The  house  finch,  Carpodacus 


Organic  Conjiection  Between  Pliysical  and  Psychical     261 

Mexicanus,  and  the  Califurnia  tuwlicc,  Pipilo  fimciis,  may  be 
specially  mentioned  in  tills  connection.  The  fact  that  do- 
mesticated song  birds,  like  the  canary,  may  be  brouglit  to 
sing  almost  perpetually  is  only  an  extreme  manifestation 
of  tendency  among  song  birds  to  sing  in  excess  of  any  strict 
utility  of  song. 

Think  of  tlie  monotonous  repetition  in  the  croaking  of 
frogs,  the  chirping  of  crickets,  the  stridulations  of  cicadas, 
and  so  on !  I  have  counted  more  than  five  hundred  con- 
secutive chirps  of  a  cricket  in  about  half  an  hour,  with 
only  a  little  variation  as  to  notes  or  intervals.  And  tliis  is 
surely  a  very  moderate  example  of  what  actually  occurs — 
as  any  one  can  easily  convince  himself  by  listening  and  count- 
ing almost  any  still  night,  almost  anywhere  where  crickets 
live.  Probably  the  chirping  of  crickets  is  employed  in  mat- 
ing. Very  well.  But  are  the  thousands  of  cliirps  uttered 
by  a  given  individual  each  niglit  for  many  nights,  the  small- 
est number  upon  which  the  species  can  survive  .^^  Even  ask- 
ing of  the  question  reveals  the  monstrosity  of  a  theory  that 
would  necessitate  an  affirmative  answer  to  it — as  strict  ad- 
herence to  the  natural  selectionist  meaning  of  utility  un- 
doubtedly would. 

In  place  of  bringing  forward  additional  instances,  which 
could  easily  be  done,  to  show  that  vocal  sounds  and  bodily 
performances  of  various  sorts  more  or  less  obviously  con- 
nected with  mating  among  higher  animals  are  produced  in 
excess  of  what  the  strict  application  of  the  rule  of  physio- 
logical economy  would  dictate,  I  shall  do  no  more  than  util- 
ize the  conclusions  of  two  investigators  who  seem  specially 
qualified  to  speak  on  the  subject,  and  assume  that  these  con- 
clusions would  receive  the  sanction  of  all  zoologists  who  have 
given  serious  attention  to  the  matter  and  have  formed  their 
judgments  unbiased  in  favor  of  any  explanatory  theory. 

The  first  of  these  investigators  is  W.  II.  Hudson,  who 
represents  a  period  a  little  antecedent  to  the  present  spc- 


262  The  Unity  of  the  Organism 

cially    critical    experimental    era.      I    quote    from    his    well- 
known  The  Naturalist  in  La  Plata,  published  in  1892:     "I 
wish  now  to  put  this  question :     What  relation  that  we  can 
see  or  imagine  to  the  passion  of  love  and  the  business  of 
courtship    have    these    dancing    and    vocal    performances    in 
nine  cases  out  of  ten?     In  such  cases,  for  instance,  as  that 
of  the  scissor-tail  tyrant-bird,  and  its  pyrotechnic  evening 
displays,  when  a  number  of  couples  leave  their  nests,  con- 
taining eggs  and  young,  to  join  in  a  wild  aerial  dance;  the 
mad   exhibitions   of   ypecahas    and   ibises,    and   the   jacanas' 
beautiful  display   of  grouped  wings ;  the  triplet  dances   of 
the  spur-winged  lapwing,  to  perform  which  two  birds  already 
mated  are  compelled  to  call  in  a  third  to  complete  the  set; 
the  harmonious  duets  of  the  oven-birds,  and  the  duets  and 
choruses  of  nearly  all  the  wood-hewers,  and  the  wing-slap- 
ping aerial  displays   of  the  whistling  widgeons ;  will  it  be 
seriously   contended  that  the   female   of  this   species  makes 
choice  of  the  male  able  to  administer  the  most  vigorous  and 
artistic   slaps?   .   .   .  There   are  many   species   in  which  the 
male,  singly  or  with  others,  practises  antics  or  sings  during 
the  love-season  before  the  female;  and  when  all  such  cases, 
or   rather  those  which  are  most   striking  and   bizarre,   are 
brought  together,  and  when  it  is  gratuitously  asserted  that 
the  females  do  choose  the  males  that  show  off  in  the  best 
manner  or  that  sing  best,  a  case  for  sexual  selection  seems 
to  be  made  out.     How  unfair  the  argument  is,  based  on  these 
carefully   selected   cases   gathered    from   all   regions    of   the 
globe,  and  often  not  properly  reported,  is  seen  when  we  turn 
from  the  book  to  Nature,  and  closely  consider  the  habits  and 
actions  of  all  the  species  inhabiting  any  one  district.     We 
see  then  that  such  cases  as  those  described  and  made  so  much 
of  in  the  'Descent  of  Man,'  and  cases  like  those  mentioned 
in   this   chapter,   are  not  essentially   different   in   character, 
but  are  manifestations  of  one  instinct,  which  appears  to  be 
almost  universal  among  the  higher  animals.     The  explana- 


Organic  Connection  Between  'Physical  and  PsycliicaJ     2(53 

tion  I  Ih'ivc  to  oft'i'j-  lios  very  iiiiicli  on  the  .surface,  .  .  .  We 
see  that  tlie  inferior  animals,  wlien  the  conditions  of  life  are 
favorable,  are  subject  to  periodical  fits  of  gladness,  affecting 
them  powerfully,  and  standing  out  in  vivid  contrast  to  their 
ordinary  tem])er.  And  we  know  what  this  fecliiiiJ-  is — this 
periodic  intense  elation  which  even  civilized  man  occasionallv 
experiences  wheji  in  perfect  health,  more  especially  when 
young.  There  are  moments  when  he  is  mad  with  joy,  when 
he  cannot  keep  still,  when  his  impulse  is  to  sing  and  shout 
aloud  and  laugh  at  nothing,  to  run  and  leap  and  exert  him- 
self in  some  extravagant  way."  ^^ 

The  reader  is  asked  to  note  what  Hudson  says  about  pick- 
ing out  such  evidence  as  will  help  the  case  for  sexual  selec- 
tion, and  saying  nothing  about  evidence  which  will  not  hel]) 
it.  Beyond  question  the  dogma  of  natural  selection,  espe- 
cially the  Weismannian  perversion  of  it,  has  flourished 
largely  on  this  sort  of  thing.  Nor  has  natural  selection  alone 
among  biological  theories  had  the  benefit  of  assorted  evi- 
dence. Indeed  the  whole  elementalistic  mode  of  interpret- 
ing living  nature  may  be  characterized  as  one  whose  doc- 
trines depend  largely  upon  "special  privilege,"  to  adopt  a 
phrase  lately  much  used  in  the  economic  world,  as  to  evi- 
dence for  their  support. 

The  other  investigator  upon  whom  we  draw  is  Prof.  Julian 
S.  Huxley,  whose  work  is  that  of  a  field  zoologist  imbued 
with  the  exacting  spirit  of  the  present  day.  Huxley's  stud- 
ies are  devoted  to  the  mating  habits  of  birds,  so  there  can 
be  no  question  that  the  activities  he  describes  are  intimately 
connected  with  reproduction.  Of  thi-  numerous  s})ecies 
dealt  with  in  the  paper  now  before  us,  we  notice  first  the 
Great  Crested  Grebe.  It  is  highly  significant  that  in  this 
species  mating  takes  place  before  the  so-called  courtship 
performances  begin,  so  this  latter  process  can  not  be  es- 
sential to  securing  a  mate.  The  female  is  "courted"  after 
she  is  got  possession  of.     The  courtship  activities  begin  soon 


264  The  Unity  of  the  Organism 

after  pairing,  two  entirely  different  sets  of  ceremonies  be- 
ing involved  in  the  activities.  One  of  these  Huxley  calls 
ceremonies  of  mutual  display,  the  other,  ceremonies  of  coi- 
tion. The  highly  elaborate  mutual  display  performances 
are  fully  described  but  can  not  be  reproduced  here.  They 
consist  in  a  variety  of  body  attitudes,  head  and  wing  and 
feather  movements,  swimmings  and  divings,  and  call-notes, 
the  w^hole  lasting  some  minutes.  Concerning  this  prelim- 
inary operation,  Huxley  writes : 

"The  most  noticeable  thing  about  all  these  ceremonies  is 
that  they  are  'self-exhausting' — they  do  not  lead  on  to  any- 
thing further.  Looked  at  from  the  physiological  point  of 
view,  they  seem  to  me  to  be  notliing  but  'expressions  of  emo- 
tion' :  the  birds  act  thus  because  they  are  impelled  to  do 
so,  because  they  enjoy  it.  Looked  at,  on  the  other  hand, 
from  the  evolutionary  point  of  view,  they  seem  to  have  been 
developed  as  a  bond  to  keep  the  pair  together."  ^^ 

Following  these  preliminaries,  the  ceremonies  of  coition 
take  place,  these  being  less  striking,  though  characteristic. 

Speaking  of  his  studies  on  the  mating  habits  of  some  of 
the  warblers,  and  referring  to  differences  of  interpretation 
between  himself  and  W.  P.  Pycraft,  another  observer  in  the 
same  field,  Huxley  writes :  "In  this,  Mr.  Pycraft  and  myself 
are,  I  tliink,  agreed;  to  both  of  us  the  'display'  of  the  male 
Warbler  is  nothing  but  a  direct  expression  of  sexual  excite- 
ment, scarcely,  if  at  all,  modified  by  Darwinian  Sexual  Se- 
lection— nothing  but  the  way  in  which  nervous  disturbance 
caused  by  sexual  excitement  happens  to  liberate  itself.  Gen- 
eral nervous  discharge  will  cause  general  muscular  contrac- 
tion;  and  something  approaching  this  is  here  seen — rapid 
hopping,  extension  and  fluttering  of  the  wings,  spreading 
of  the  tail,  bristling  up  of  the  fea.thers  on  head  and 
throat,  and  utterance  of  a  series  of  quick  sounds.  This  ex- 
presses a  condition  of  readiness  to  pair,  and  doubtless  to 
the   female   comes    to   be   a   symbol   of   the   act   of   pairing. 


Organic  Connection  Between  Physical  and  Psijchical     ^65 

Hence,  as  far  as  the  female  is  concerned,  the  act  of  pairing 
has  come  to  dej)cnd  upon  this  stimuhis  (acting  of  course  on 
a  suitable  internal  j)hvsiological  state).  This  is  no  more 
strange  in  the  bird  than  it  is  that  in  ourselves  thoughts  and 
emotions  of  love  well  up  at  the  sight  of  some  tangible  ob- 
ject connected  with  the  beloved,"  ^'^ 

But  it  is  in  the  sex  function  itself  that  the  tendency  to 
overdo  manifests  itself  with  greatest  force.  In  fact,  the  fa- 
miliar and  ominous  expression  "sexual  excesses"  as  applied 
to  the  human  animal,  indicates  very  truthfully  what  is  be- 
fore us.  The  whole  phenomenon  of  competing  and  fighting 
among  the  males  of  all  higher  animals  for  possession  of  the 
females,  with  its  momentous  consequences  in  dozens  of  ways, 
may  truly  be  said  to  rest  back  on  the  excessiveness  of  the 
sex  impulse  and  instinct.  Since  as  a  general  rule  the  males 
and  females  of  animal  species  are  approximately  equal  in 
numbers,  pairing  off  two  by  two,  after  the  manner  of  the 
population  of  Noah's  ark,  might  occasion  but  little  and  mild 
competition  could  each  male  and  each  female  be  satisfied 
with  one  mate,  in  accordance  with  the  allotment  which  the 
numerical  equality  would  make.  And  the  pertinent  question 
may  be  raised  in  passing,  would  not  such  a  mode  of  pairing 
secure  the  perpetuation  of  the  species  quite  as  well  as,  pos- 
sibly better  than,  the  method  which  is  so  largely  in  vogue  .'^ 

Highly  suggestive  seems  to  me  in  this  connection,  observa- 
tions I  have  recently  been  able  to  make  on  the  mating  hab- 
its of  one  of  the  California  "surf  perches"  {Cymatogastcr 
aggregatus).  This  is  one  of  the  numerous  viviparous  bony 
fishes  peculiar  to  our  coast.  The  species  under  attention 
lives  quite  normally,  as  far  as  one  can  see,  in  the  aquaria  of 
the  Scripps  Institution;  so  what  may  be  assumed  to  be  its 
typical  habits  can  be  observed  continuously. 

Strict  monogamy  appears  to  prevail  in  the  species.  At 
least  this  is  true  with  the  specimens — three  males  and  four 
females  under  observation,  and  so  far  as  a  particular  breed- 


266  The  Unit!/  of  the  Organism 

ing  period  is  concerned.  Each  male  begins  his  attentions 
while  his  fiancee,  so  to  speak,  is  heavily  gravid  from  the  pre- 
vious mating  (when  and  how  accomplished  we  unfortunately 
know  nothing  about  beyond  the  fact  that  it  must  have  been 
before  the  individuals  under  observation  were  brought  to 
the  aquarium  from  the  sea,  about  six  weeks  before  the  mat- 
ing began.) 

In  the  case  of  one  pair,  the  amours  of  the  male  continued 
more  than  two  weeks,  the  first  few  days  of  which  were 
before  the  family  of  young  began  to  be  born,  the  period  of 
parturition  extending  over  three  days.  Although  there 
was  no  indication  on  the  part  of  the  other  males  of  inten- 
tions or  even  desires  toward  the  spouse  (as  she  may  now 
be  called)  of  this  male,  he  was  quite  pugnacious,  directing  his 
seemingly  unnecessary  operations  against  the  other  females 
as  well  as  against  the  other  males.  It  should  be  said,  how- 
ever, that  his  antipathies  were  considerably  greater  against 
the  males  than  against  the  other  females.  The  other  two 
males  took  partners  after  much  the  same  fashion ;  but  since 
both  of  these  were  somewhat  smaller,  and  fully  acknowl- 
edged the  over-lordship  of  the  one  singled  out  in  our  account, 
their  performances  were  less  clear  cut. 

Specially  noteworthy  is  the  character  of  the  amours  of 
the  male,  which  alone  or  almost  alone,  seems  to  take  an  in- 
terest in  the  performance.  No  contact,  or  at  least  only  the 
slightest,  of  the  male  with  the  female  was  seen  though  the 
fish  were  under  observation  much  of  the  time.  A  peculiar 
downward  darting  of  the  male  first  on  one  side  then  on  the 
other  of  the  female,  close  to  her  but  not  quite  touching  her, 
was  one  of  the  favorite  manoeuvers.  But  various  rapid 
circlings  about,  up  and  down,  head-on  and  tail-on,  over  and 
under,  and  in  nearly  all  possible  ways,  may  be  witnessed. 

The  full  meaning  of  this  monogamic  (temporarily  so,  at 
least),  largely  non-tactual  type  of  mating  we  do  not  know 
partly  because  we  have  not  yet  all  the  facts ;  but  I  suspect 


Organic  Connection  Between  Physical  and  Pstjchical     267 

it  to  be  important.  But  this  much  is  clear  as  to  its  bear- 
ing upon  the  point  uppermost  in  tliis  discussion:  There  is 
an  excessiveness  of  activity  in  a  variety  of  ways,  particuhirly 
in  the  driving  of  other  females,  the  presence  of  which  in  the 
vicinity  of  the  mate  is  merely  incidental  and  utterly  harm- 
less. 

Obviously  it  is  tlie  demand,  instinctive  or  organic  or  both, 
for  more  sexual  gratification  than  the  natural  numerical 
scheme  of  the  two  sexes  provides,  and  the  actual  necessities 
of  race  perpetuation  deman.d,  wlilch  is  largely  responsible 
for  the  contests  to  secure  mates,  so  characteristic  of  all 
higher  animals.  The  bull  fur  seal  must  have  forty  or  fifty 
mates,  instead  of  the  one  which  tlie  numerical  equality  of 
the  two  sexes  would  naturally  give  him ;  hence  the  fierce  com- 
bats among  the  males,  with  the  result  that  a  great  majority 
of  the  whole  male  population  at  any  one  time  is  forced  to 
remain  outside  the  "harems"  during  the  mating  season.  And 
some  such  eliminative  process  must  occur  in  all  species  where 
the  sexes  are  about  equal  in  numbers,  and  where  promiscuity 
in  pairing  is  practised. 

Nor  are  the  injuries  and  disasters  which  may  result  from 
the  driving  power  of  the  sex-impulse  restricted  to  compet- 
ing individuals  of  the  same  sex.  The  mates  sought  after  not 
infrequently  suffer  seriously  from  the  excesses  of  the  seek- 
ing males,  the  females  being  usually  more  passive  and  hence 
the  more  liable  to  injury  in  this  way.  Thus,  J.  S.  Huxley 
has  lately  told  of  the  exliaustion  and  actual  death  of  the 
female  mallard  duck  from  being  repeatedly  "tread"  by  the 
males,  the  same  and  different  individual  males  participating 
in  the  strangely  destructive  performance. 

Finally,  the  individual  itself  is  not  safe  from  self-injury 
through  its  own  sex  impulses.  Some  of  the  forms  which  this 
sort  of  thing  may  take  in  tlie  human  species  are  too  famil- 
iar, too  disastrous  and  too  repugnant  to  need  illustration 
in  proof  of  their  reality.      That  they  occur  also  more  or  less 


268  The  Unity  of  the  Organism 

among  animals  is  well  known  to  all  who  have  had  consid- 
erable experience  with  domestic  animals. 

Excessive  activity  in  connection  with  the  alimentary  func- 
tion must  now  be  glanced  at.  That  there  is  no  nice  quanti- 
tative balance  between  the  food  necessities  of  the  animal  and 
the  food  gathering  instincts  and  impulses  and  efforts  on  the 
basis  of  the  principle  of  natural  economy  and  parsimony, 
is  shown  conclusively  it  would  seem  by  many  animals  which 
have  the  storing  habit.  The  honey  bee  is  an  example  of 
this  among  insects.  Given  a  sufficient  supply  of  flowers  to 
work  on,  in  the  wild  state  these  bees  seem  always  to  store 
away  more  food  material  than  they  consume. 

The  extent  of  their  honey-making  is  limited  rather  by 
the  raw  material  available  and  by  their  own  restricted  phys- 
ical powers  than  by  their  nutritional  needs.  This  is  the  im- 
pression I  have  from  my  observations  on  wild  and  tame  bees 
and  I  find  it  to  coincide  with  that  of  other  naturalists  whose 
opportunity  for  obsei-Aang  wild  bees  has  been  much  greater 
than  mine.  For  example  my  esteemed  naturalist  friend,  Mr. 
Frank  Stephens  of  San  Diego,  California,  reminds  me  that 
the  view  is  confirmed  by  the  fact  that  in  "bee  trees"  a  por- 
tion of  the  comb  containing  honey  is  not  infrequently  black 
and  shows  signs  of  being  old. 

Darwin  made  quite  a  point,  it  may  be  recalled,  of  the 
economy  in  some  aspects  of  the  bee's  work.  "The  comb  of 
the  hive-bee,"  he  says,  "as  far  as  we  can  see,  is  absolutely 
perfect  in  economising  labour  and  wax."  {Cell-Making  In- 
stinct of  the  Hive-Bee,  in  The  Origin  of  Species.)  ^*  But  a 
thoroughly  economic  adjustment  between  different  parts  of 
a  given  complicated  operation,  and  economy  of  the  opera- 
tion as  a  whole,  are  very  different. 

As  an  instance  of  excessive  repetition  in  the  food-getting 
activities  among  the  insects,  the  following  from  Fabre  may 
be  taken  as  fairly  typical.  A  solitary  wasp  of  the  genus 
Sphex  captures  and  slays  a  locust,  but  instead  of  using  it 


Organic  Connection  Between  Physical  and  Psychical     269 

at  once  for  food,  or  of  taking  it  directly  into  her  home,  she 
sometimes  leaves  it  on  the  road,  and  nins  to  her  home,  even 
though  this  is  threatened  by  no  danger.  Then  after  a  time 
she  returns  to  the  game.  Tliis  going-and-coming  may  be 
performed  repeatedly  before  the  carcass  is  finally  taken  into 
the  dwelling.  If  by  chance  the  game  is  removed  during  the 
absence  of  the  wasp,  the  wasp  returns  to  the  spot  where  her 
load  was  left,  but,  not  finding  it,  she,  nevertheless,  keeps  up 
the  going-and-coming  for  some  time.  The  first  back-and- 
forth  journey  from  game  to  dwelling  is  explicable,  Fabre 
shows.  "But  what  is  the  use  of  the  other  visits,  repeated 
so  speedily  one  after  another.'"'  Fabre  inquires.^ ^  Something 
like  this  almost  every  one  must  have  seen,  who  has  watched  in- 
sects at  all. 

I  am  quite  certain  that  the  acorn  storing  habit  of  the 
California  woodpecker,  Melanerpes  formicivorus  bairdi,  is 
quite  beyond  any  use  the  bird  makes  of  the  acorns.  In  the 
first  place,  despite  much  discussion  of  the  question  whether 
the  acorns  are  used  at  all,  and  if  so  how,  the  case  is  by  no 
means  clear.  But  the  point  I  particularly  wish  to  make  is 
that  whatever  use,  if  any,  the  birds  make  of  the  acorns, 
whether  as  food  directly  or  as  culture  media  for  worms  or 
insects,  these  in  turn  to  be  eaten  by  the  birds,  they  store  up 
many  more  than  they  utilize.  This  seems  to  me  highly  prob- 
able from  the  fact,  which  I  have  ascertained  by  numerous 
examinations  at  different  places  and  times,  that  many  holes 
contain  dried  up  and  wasted  acorns  which  show  no  signs 
of  having  been  picked  at  or  otherwise  moved  after  they  were 
inserted  into  the  holes.  Furthermore,  the  great  extent  of 
the  hole-drilling  and  filling  in  itself  seems  to  exceed  the 
bounds  of  necessity,  especially  in  view  of  the  certainty  that 
the  bird's  chief  food  supply  is  from  quite  another  source.  A 
pine  log  fifty  feet  long  and  one  hundred  thirty-six  inches  in 
girth  at  the  middle,  which  I  found  in  the  San  Jacinto  Moun- 
tains, contained   on   a  fairly   careful  estimate  31,800  holes. 


270  The  Unity  of  the  Organism 

many  of  them  containing  acorns. 

But  even  were  it  certain  that  the  acorns  are  utilized  in  any 
manner  and  to  some  extent  in  connection  with  the  feeding 
function,  there  are  still  other  evidences  than  that  just  ad- 
duced of  the  imperfect  and  excessive  operation  of  the  acorn- 
storing  instinct.  As  is  well-  known,  the  bird  sometimes  ex- 
tends its  drilling  operations  to  wooden  buildings  to  the  ex- 
tent of  making  itself  a  great  nuisance.  I  have  seen  a  case 
where  the  birds  had  pierced  the  rustic  of  an  uninhabited 
house,  so  that  when  the  acorns  were  inserted,  instead  of 
filling  the  puncture  as  they  would  fill  holes  in  a  tree,  they 
would  drop  down  into  the  space  between  the  rustic  and  the 
inner  wall.  Apparently  the  failure  to  stop  the  hole,  and 
failure  also  to  perceive  why,  or  to  recognize  that  the  hole 
could  not  thus  be  stopped,  "fooled"  the  birds  into  putting 
one  acorn  after  another  into  the  same  hole,  endlessly  almost, 
judging  by  the  great  quantity  of  nuts  piled  up  at  the  bot- 
tom of  the  space. 

While  the  storing  habit  of  the  California  woodpecker  is 
undoubtedly  exceptional  as  to  extent,  it  is  by  no  means 
wholly  unique.  At  least  one  species  of  blue- jay  (Cyanocitta 
cristata)  has  much  the  same  habit,  in  the  opinion  of  most 
ornithologists  who  have  studied  the  habits  of  the  bird.  An 
experienced  naturalist,  E.  H.  Forbush,  has  recently  said 
concerning  Mark  Twain's  "Baker's  Blue  Jay  Yarn,"  in  A 
Tramp  Abroad,  "All  of  this  is  not  merely  amusing;  it  is  good 
ornithology  in  so  far  as  it  reports  the  way  a  Jay  acts."  ^^ 
This  story,  it  may  be  said  for  the  benefit  of  any  reader  so 
unfortunate  as  not  to  know  it,  turns  upon  the  performance 
of  a  jay  similar  to  that  narrated  above  about  the  California 
woodpecker,  the  acorns,  and  the  old  house. 

The  habit  of  the  shrikes  (genus  Lanius)  of  impaling  their 
victims  and  leaving  them,  almost  certainly  operates  more  or 
less  independently  of,  and  often  in  excess  of,  the  food  re- 
quirements of  the  birds.     ''My  observations,"  says  Forbush, 


Organic  Connection  Bet-ween  Physical  and  Psijchical     5^71 

"have  led  nie  to  believe  that  it  rarely  returns  to  eat  what  it 
has  thus  cached,  unless  driven  to  do  so  by  liunfrer  resulting 
from  adverse  fortunes  of  tlie  chase."  ^'^ 

Nor  is  tliere  much  if  any  question  that  something  of  the 
same  sort  occurs  among-  mammals  wliich  have  the  food  stor- 
ing liabit.  p].  T.  Seton  quotes  the  following  from  Dr.  John 
Wriglit  concerning  the  big  eastern  chipmunk  {Tamias  strin- 
tus  grisens)  :  "It  is  a  most  provident  little  creature,  con- 
tinuing to  add  to  its  winter  store,  if  food  is  abundant,  until 
driven  in  by  the  severity  of  the  frost.  Indeed,  it  seems  not 
to  know  when  it  has  enough,  if  we  may  judge  by  the  surplus 
left  in  the  spring,  being  sometimes  a  peck  of  corn  or  nuts 
for  a  single  squirrel."  ^^  There  are  many  other  statements 
by  the  best  authorities,  especially  concerning  numerous  spe- 
cies of  mice,  which  strongly  suggest  a  like  superabundance 
of  storing  activities.  But  for  the  rest  I  w^ill  mention  a  case 
that  has  come  to  my  own  notice. 

I  am  indebted  to  jMr.  Frank  Stephens  for  information 
about  and  the  opportunity  to  witness  to  some  extent  for 
myself  the  operations  of  the  storing  instinct  and  feeding 
habits  of  the  Antelope  Ground-Squirrel  {Amnios pcrinophi- 
lus  leucurus).  This  chipmunk-like  little  squirrel  proves  to 
be  so  readily  domesticable  that  it  becomes  almost  as  famihar 
a  household  member,  at  least  for  ]\Ir.  Stephens'  household, 
as  a  domestic  cat.  Altliougli  an  account  of  the  habits  of  the 
single  individual  in  Mr.  Stephens'  possession  can  not  yet  be 
told  fully  by  a  long  ways,  a  few  points  of  much  interest 
for  the  present  discussion   are  ])ositive  enough. 

In  tlie  first  ])lace  the  genuinely  instinctive  character  of 
the  storing  habit  is  establislied  by  the  fact  tliat  althougli  the 
specimen  under  observation  was  taken  soon  after  birth,  and 
has  lived  all  its  life  in  complete  isolation  from  parents  and 
all  its  kind  and  has  been  furnished  artifii-ially  with  an 
abundance  of  food,  its  storing  operations  are  carried  o?i 
constantly  and  almost  as  perfectly,  so  far  as  one  can  .jiulgv. 


272  The  Unity  of  the  Organism 

as  though  it  were  living  in  the  natural  state.  This  fact  in 
itself  is  evidence  that  the  instinct  is  not  determined  solely 
by  immediate  needs  of  the  individual.  But  much  more 
convincing  evidence  furnished  by  this  case  to  this  effect  is  in 
the  particular  way  the  instinct  works.  For  example,  this 
species  possesses  cheek  pouches  for  carrying  food  as  do  so 
many  rodents  which  have  the  storing  habit.  When  nuts, 
grain,  etc.,  are  presented  to  the  animal  she  very  rarely  eats 
them  immediately  even  though  manifestly  hungry,  but  carries 
them  away  to  some  distance,  one  at  a  time ;  going  back  and 
forth  and  placing  the  articles  in  her  two  pouches  till  these 
are  quite  full.  And  these  little  pre-storage  journeys,  as 
they  may  be  called,  are  often  definite  in  character.  At  any 
given  time  they  end  at  nearly  the  same  spot,  and  the  animal 
takes  nearly  the  same  position  while  the  article  is  being  pre- 
pared for  and  inserted  into  the  pouches.  This  is  clearly 
the  typical  procedure  in  filling  the  pouches,  though  it  is 
varied  considerably  from  time  to  time. 

As  to  what  follows  the  pouch-filling  there  is  considerable 
variation — noraially  so  it  appears.  In  case  the  animal  is 
hungry  she  may  quietly  extract  the  nuts  from  the  pouches 
and  eat  them.  Or  she  may  run  about  for  some  time  with 
her  cheeks  bulging  full.  Or  she  may  take  her  load  off  some- 
where and  lay  it  away  either  in  some  cache  previously  es- 
tablished or  in  a  new  one.  The  cache  may  be  in  a  bed  of  sand 
if  this  is  at  hand ;  or  it  may  be  in  or  under  some  old  garment 
or  piece  of  cloth  or  paper  which  the  surroundings  may 
present. 

An  especially  interesting  fact  noticed  by  Mr.  Stephens 
is  the  tendency  shown  on  the  part  of  this  squirrel  to  carry 
the  articles  to  as  distant  a  place  from  where  it  gets  them 
as  can  well  be  reached. 

On  the  whole  there  is  no  doubt  that  we  have  here  a  var- 
iedly illustrative  example  of  activity  over  and  above  need 
in  the  operation  of  an  instinct. 


Organic  Connection  Between  Physical  and  Psjjchical     ^Ti) 

This  bare  touch,  so  far  as  instances  are  concerned,  of 
overactivity  in  connection  with  reflexes,  and  especially  with 
instincts  wliich  are  on  tlie  wliole  useful,  leads  naturally  to 
the  great  field  of  animal  play.  Space  limitations  ])rohibit 
us  from  taking  more  than  a  bird's  eye  view  of  this  field. 
Fortunately,  however,  even  such  a  view  can  be  quite  effective 
for  our  purpose  because  of  the  well-known  work  of  Karl 
Groos,  The  Ploy  of  Animals.  Our  sole  purpose  here,  as 
in  the  rest  of  this  discussion,  is  to  answer  the  question 
whether  animals  do  or  do  not  carry  their  activities  which  on 
the  whole  are  fundamental  to  their  existence  be3'ond  what 
is  necessary  for  their  own  individual  requirements.  With 
Groos's  explanatory  theory  of  play  we  are  concerned  only 
so  far  as  it  involves  the  question  of  fact  upon  which  our 
present  interest  centers.  That  most  if  not  all  animal  activ- 
ity which  can  rightfully  be  called  y)lay,  and  which  is  not 
intelligent,  is  instinctive,  we  believe  Groos  has  conclusively 
shown.  The  explanation  adopted  by  Spencer  and  others 
that  play  is  the  useless  imitation  by  young  animals  of  useful 
activities  performed  by  their  seniors,  the  imitative  acts  be- 
ing useless  because  merely  the  overflow  of  "surplus  energy," 
is  certainly  inadequate,  as  Groos  has  insisted.  That  animals 
constantly  go  through  perfomiances  playfully  which  they 
have  had  no  chance  to  see  or  to  have  otherwise  impressed 
upon  them  from  without,  is  as  certain  as  that  they  constantly 
perform  useful  acts  in  this  way. 

It  consequently  results  that  a  source  of  energy  for  play, 
that  is,  for  actions  which  are  not  immediately  essential  to 
the  existence  of  the  organism,  must  be  an  endowment  of  tin 
organism  no  less  certainly  than  that  a  source  of  energy  mu>f 
exist  for  actions  which  are  essential  to  its  existence.  So 
Groos's  statement:  "A  condition  of  sur})lus  energy  still  ap- 
pears as  the  conditio  sin^  qua  noii  that  pennits  the  force 
of  the  instincts  to  be  so  augmented  that  finally,  when  a  real 
occasion  for  their  use  is  wanting,  they  form  their  own  mo- 


274  Tine  Unity  of  the  Organism 

tive,  and  so  permit  indulgence  in  merely  sportive  acts,"  ^^ 
becomes  a  statement  of  fact  if  by  "surplus  energy"  we 
understand  energy  available  for,  and  upon  occasion  used  for, 
acts  wliich  are  not  indispensable  to  the  existence  of  the  in- 
dividual. 

The  quantity  and  generality  of  play  performed  by  animals 
may  be  taken  as  one  important  measure  of  the  extent  of  the 
energy  possessed  over  and  above  what  is  essential  for  their 
normal  individual  existences,  and  this  without  reference  to 
whether  or  not  the  play  may  be  useful  as  a  preparation  for 
future  essential  activities,  or  for  recreation  only.  The  fact 
can  hardly.be  too  much  insisted  upon  that  ulterior  useful- 
ness of  the  organism's  acts,  whether  to  the  species  generally, 
to  offspring,  or  to  the  individual's  own  future,  cannot  pos- 
sibh^  be  a  sufficient  explanation  of  the  energy  immediately  re- 
quired for  the  act  itself.  Even  though  an  animal  does  noth- 
ing whatever  except  by  reason  of  its  hereditary  endowments, 
or  in  the  interest  of  its  offspring;  and  though  the  real  pur- 
pose of  much  that  it  does  looks  to  its  own  future,  it  must 
nevertheless  continue  to  eat,  digest  and  assimilate,  and 
breathe.  The  subdivision  of  biology  which  has  come  to  be 
known  as  physiology  has  for  its  distinctive  task  exactly 
that  of  studying  the  present  activities  of  the  organism.  With 
the  organism's  past,  whether  individual  or  racial,  and  with 
its  future,  whether  individual  or  racial,  physiology  can  be 
concerned  only  indirectly. 

Summary    of    Orgcmismal    Character    of    All    Suhrational 

Psychic  Life 

Having  now  examined  broadly  though  far  from  exhaus- 
tively the  psychic  life  of  the  animal  in  each  of  its  most  ob- 
vious phases,  the  highest  rational  phase,  the  emotional  phase, 
the  instinctive  phase,  and  the  reflex  phase  (in  which  tropisms 
are  included)  for  purely  descriptive  and  classificatory  pur- 


Organic  Connection  Between  Phi/sical  and  Psijchical     275 

poses,  let  us  briefly  summarize  wlint  we  have  learned. 

In  each  phase  we  have  found  the  oryunism,  living,  whole 
and  normal,  indispensable  to  a  comprehension  of  the  phe- 
nomena examined.  Or,  expressed  in  a  different  way,  we  have 
found  it  possible  in  each  phase  to  reach  only  a  very  imperfect 
understanding  of  the  phenomena  by  referring  tliem  to  the 
elements  which  can  be  discovered  in  them.  For  example,  the 
theory  of  association  of  ideas  is  inadequate  to  explain  ra- 
tional life,  in  such  manifestations  as  apperception  and  mental 
initiative  and  creativeness. 

In  the  emotional  phase,  in  such  emotions  as  fear,  rage  and 
sex  passion,  not  only  does  cursory  observation  recognize 
the  involvement  of  a  large  part  of  the  organism,  but  physi- 
ological investigation  is  able  greatly  to  extend  our  recogni- 
tion of  this  involvement  by  showing  how  the  nervous  system 
in  its  cerebro-spinal  and  its  autonomic  divisions,  the  cir- 
culatory, the  alimentary,  and  the  internal-secretory  sys- 
tems, are  essentially  and  reciprocally  involved. 

As  to  the  organismal  character  of  psychic  life  in  the 
phase  of  instinct,  it  suffices  to  recall  that  one  of  the  most 
widely  accepted  criteria  for  differentiating  instinctive  from 
reflex  activities  is  that  the  former  involve  the  organism  as  a 
unity,  a  whole,  while  reflexes,  according  to  this  criterion, 
pertain  only  to  limited  portions  of  the  organism.  '*An 
instinct  is  a  more  or  less  complicated  activity  manifested  by 
an  organism  which  is  acting,  first,  as  a  whole  rather  than  as 
a  part."  -"^  To  this  statement  of  the  matter  may  be  added 
that  when  the  instinctive  act  is  in  the  interest  of  the  indi- 
vidual performing  it,  the  act  is  not  only  by  but  for  the  in- 
dividual as  a  whole. 

As  to  the  reflex  phase  (if  that  is  to  be  reckoned  as  psy- 
chic) the  organismal  nature  of  tropisms  has  liad  so  large 
a  place  in  our  discussion  that  surely  no  more  nerd  l)e  done  in 
this  summary  than  to  remind  the  reader  of  our  discussion  of 
tropisms.      And   even   reflexes   of   a  simpler  fDiifi   than    tlie 


276  The  Unity  of  the  Organism 

tropisms — indeed  the  abstract  conception  of  the  "simple 
reflex,"  though  not,  perhaps,  involving  the  conception  of  the 
organism  as  a  whole,  yet  is  not  comprehensible  on  elemen- 
talistic  principles,  as  our  examination  of  Sherrington's  in- 
vestigations revealed.  And  such  phenomena  as  those  of  the 
spreading  and  compounding  of  reflexes  are  quite  incompre- 
hensible except  on  the  organismal  principle,  even  though  the 
whole  organism  may  not  be  involved,  observably  at  least,  in 
particular  reflex  acts. 

Specificity  of  Siihrational  Psychic  Life 

The  concluding  section  of  this  descriptive  chapter  on 
psychic  integratedness  must  be  devoted  to  the  specificity, 
not  to  say  individualit}^,  of  animal  behavior  in  all  its  phases. 
The  vast  body  of  trustworthy  detailed  knowledge  now  in  our 
possession  justifies,  I  am  quite  sure,  the  following  general- 
ized statement  under  this  head:  It  is  exactly  on  the  psychic 
side  of  animal  life,  psychic  being  taken  in  the  broadest  sense, 
that  animals  are  most  differentiated  from  one  another,  both 
as  to  individuals  and  as  to  species. 

Taxonomic  zoology  is  based  almost  entirely  on  structural 
attributes  of  animals.  This  results  from  reasons  that  are 
obvious,  speaking  generally,  and  constitutes  a  justification 
of  the  fact  from  a  practical  standpoint.  Nevertheless  the 
purely  practical  advantages  of  the  classificatory  systems  as 
they  have  been  built  up  have  been,  and  are,  gained  at  the 
expense  of  several  rather  serious  disadvantages.  One  of 
these  is,  as  advance  of  knowledge  leads  us  to  realize,  that  our 
well-nigh  exclusive  attention  to  structural  diff^erences  and 
likenesses  has  tended  strongly  to  divert  attention  from  func- 
tional differences  and  likenesses.  It  is  of  fundamental  impor- 
tance for  a  truly  comprehensive  science  of  organic  beings, 
that  is,  for  a  philosophical  biology,  to  regard  our  synoptic 
classifications  not  as  a  final  result  of  knowledge-getting,  but 


Organic  Connection  Between  Physical  and  Psijchical     277 

rather  as  a  life-sized  sketch,  as  one  may  say,  of  the  whole 
living  world,  to  facilitate  the  gigantic  task  of  completing 
the  picture  through  the  cooperation  of  numberless  artists, 
the  completion  to  be  accomplished  by  filling  in  the  sketcii 
with  the  entire  round  of  attributes,  structural  and  func- 
tional, presented  by  the  natural  lives  of  organisms.  I  have 
dwelt  somewhat  at  length  on  this  matter  elsewhere,*  and  can 
refer  to  it  here  only  as  a  background  for  what  I  wish  to  say 
about  psychical  specificity. 

Two  extracts  must  suffice.  "No  biological  phenomenon 
is  adequately  interpreted  or  dealt  with  experimentally,  until 
it  has  been  considered  with  reference  to  the  place  that  the 
organism  to  which  it  pertains  holds  in  the  system  of  classi- 
fication." And  further:  "What  I  affirm  is  that  the  inductive 
evidence  has  now  gone  so  far  toward  proving  every  sharply 
differentiated  species  to  contain  some  differentia  in  all  the 
main  provinces  of  their  structure  and  function,  that  to  as- 
sume the  absence  of  such  differentia  in  any  given  case  is 
unwarranted."  ^^ 

I  want  to  utilize  these  earlier  general  statements  about 
organic  specificity,  as  a  basis  on  which  to  rest  a  generaliza- 
tion concerning  the  specificity  of  psychic  attributes.  So 
enormous  is  the  observational  data  available  for  illustration 
here,  that  in  lieu  of  presenting  any  of  them  I  am  going  to 
state  in  a  wholly  dogmatic  fashion  the  generalization  toward 
which  we  are  certainly  being  led  by  modern  crucial  researches 
on  animal  behavior.  Let  us  imagine  ourselves  possessed  of  an 
approximately  exhaustive  descriptive  knowledge  of  the  be- 
havior of  the  whole  animal  world,  tliis  knowledge  Ix'ing,  liow- 
ever,  unaccompanied  by  any  knowledge  whatever  of  the  cor- 
poreal nature  of  the  animals.  This  behavior-knowledge 
would  fall  naturally  into  categories  larger  and  smaller,  the 

*  The  Place  of  Description,  Definiti<in  and  Classification  in  Philoso- 
phical Biology,  in  The  Higher  Usefulness  of  Science  (Boston,  1918); 
also  The  Scientific  Monthly,  November,  1916. 


278  Tlie  Unity  of  the  Organism 

smaller  ones  being  for  the  most  part  subdivisions  under  the 
larger.  Then  let  us  imagine  this  system  of  behavior-knowl- 
edge compared  exhaustively  with  a  later-acquired,  equally 
exhaustive  knowledge  of  the  corporeal  nature  of  all  animals. 
The  two  systems  would  be  found  to  match  each  other  very 
nearly  as  closely  as  though  the  two  had  been  worked  out  to- 
gether, much  as  they  are  being  actually  elaborated  by  struc- 
tural and  functional  zoology.  In  other  words,  the  species, 
genera,  orders,  and  so  on,  of  animals  are  differentiated  from 
one  another  and  coordinated  with  one  another  by  their  "be- 
havior," that  is,  by  their  whole  round  of  psycliical  and  reflex 
attributes,  much  as  they  are  by  their  corporeal  attributes. 

The  inductive  evidence  for  such  a  generalization  is  being 
produced  at  the  present  time  by  three  quite  definitely  marked- 
off  kinds  of  research.  These  may  be  designated  as  ( 1 )  quali- 
tative field  researches,  (2)  laboratory  experimental  re- 
searches, and  (3)  quantitative  field  researches,  the  definitely 
quantitative  method  being  statistical. 

The  first-mentioned  class  of  investigations  is  typified  by 
the  earlier  field  zoologists,  whose  aim  was  to  learn,  as  ex- 
haustively as  possible  in  a  purely  qualitative  way,  the  habits 
of  animals  in  nature.  Workers  of  this  class  are  the  typical 
zoological  naturalists  of  the  history  of  animal  biology. 
Aristotle,  Conrad  Gesner,  John  Ray,  Charles  Linne,  P.  S. 
Pallas,  Gilbert  White,  J.  J.  Audubon,  J.  H.  Fabre,  A.  R. 
Wallace  and  A.  Forel  may  be  named  as  conspicuous  examples 
of  pre-modern  members  of  this  class;  and  Charles  Darwin 
stands  out  sharply  as  a  representative  of  it,  but  as  a  transi- 
tion to  the  modern  period,  the  transitional  character  of 
Darwin  being  seen  not  only  in  the  doctrines  he  proposed  but 
as  well  in  his  intimate  combination  of  the  experimental 
method  with  the  older  method  of  obsei-vation. 

By  the  modern  period  of  research  in  field  zoology  I  mean 
the  period  during  which,  while  the  natural  history  point  of 
view  and  attitude  are  retained,  the  critical  rigor  of  modern 


Organic  Connection  Between  Pli/jsiral  and   I'sz/chiral     !27<) 

science  generally  is  practiced,  and  experimentation  in  one 
form  or  another  is  employed  as  a  supplementary  agency 
wherever  and  whenever  possible.  One  of  the  best  examples 
of  this  type  of  zoological  research  and  writing  is  Ants,  by 
W.  M.  Wheeler.  But  a  considerable  portion,  and  fortunate- 
ly an  increasing  portion,  of  experimental  research  in  animal 
behavior  is  being  done  quite  in  the  spirit  of  field  zoology. 
The  work  of  11.  M.  Yerkes  deserves  mention  as  perhaps  the 
most  definitely  purposed  and  executed  combination  of  the 
field  and  experimental  methods  for  investigating  the  behavior 
of  mammals  and  birds,  that  has  yet  been  made.  But  much 
of  the  research  recently  named  animal  ecology  tends  strong- 
ly toward  rigor  in  field  investigation.  This  kind  of  study  is 
specially  adapted  to  bring  out  the  specific  nature  of  behav- 
ior, since  the  group  of  organisms,  species,  etc.,  as  a  whole, 
occupies  a  central  place  in  the  student's  interest,  so  that  if 
behavior  is  attended  to  at  all  its  difi*erential  features  are 
likely  to  receive  attention  along  with  the  differential  struc- 
tural features. 

(2)  Laboratory  experimentation  on  animal  behavior  has, 
as  previousl3^  indicated,  been  prosecuted  more  intensely  and 
widely  in  the  modern  period  than  either  of  the  other  classes 
of  investigation.  In  fact,  it  may  be  said  to  be  distinctive  of 
the  period,  and  to  have  set  the  standard  as  regards  rigor  for 
the  other  types  of  investigation.  From  its  very  nature,  how- 
ever, it  is  not  calculated  to  bring  the  specificity  of  behavior 
to  a  central  place  in  the  student's  interest.  Singling  out 
as  it  does  one  or  a  few  attributes  at  a  time  as  they  are  ex- 
hibited by  one  or  a  few  individuals  of  one  or  a  few  species, 
breadth  and  penetration  of  comparison  are  liable  to  be  sacri- 
ficed. This  kind  of  research  tends  to  be  extremely  particu- 
laristic in  every  way.  Nevertheless,  painstaking  and  judi- 
cious workers,  like  P'.nglemann,  Forel,  Binet,  Wasmann, 
Romanes,  C.  Lloyd  Morgan,  Verworn,  Jennings,  Loch, 
Holmes,  and  Parker,  generally  state  what  species  their  in- 


280  The  Unity  of  the  Organism 

vestigations  have  dealt  with,  so  a  reader  interested  in  the 
question  of  specificity  can  usually  detect  evidences  of  dif- 
ferences in  the  behavior  of  different  species,  even  though  the 
investigator  himself  was  obviously  little  interested  in  the 
subject,  and  so  took  no  pains  to  bring  out  such  evidence. 

Indeed,  the  fact  that  species-differences  in  behavior  can  so 
frequently  be  recognized  in  descriptions  even  though  the 
writer's  general  attitude  may,  if  anything,  militate  against 
the  disclosure  of  the  differences,  is  rather  strong  evidence 
of  the  general  prevalence  of  such  differences. 

(3)  Although  statistical  investigation  of  animal  behavior 
has  been  much  less  prosecuted  than  has  either  of  the  other 
types  it  is,  nevertheless,  within  the  limits  of  its  availability, 
a  very  valuable  method  for  revealing  species  differentia,  its 
efficacy  consisting  in  the  fact  that  species  may  be  compared 
with  reference  to  different  behavior  traits  taken  one  by  one, 
and  on  the  basis  of  quantitative  data  covering  considerable 
samples  of  whole  populations.  The  method  is  specially  ap- 
plicable to  the  minute  floating  life  of  the  seas  and  lakes, 
known  as  plankton,  and  is  being  much  employed  to  this  end 
at  the  Scripps  Institution  for  Biological  Research.  It  can- 
not be  described  in  detail  here,  but  consists  essentially,  as 
employed  at  this  Institution,  in  collecting  great  quantities 
of  organisms  by  agencies  as  nearly  quantitatively  constant 
and  accurate  as  possible,  in  counting  the  organisms  thus 
secured,  and  in  correlating  the  biological  values  thus  ob- 
tained with  quantitative  studies  on  the  physical  environment 
of  the  organisms,  these  environmental  determinations  being 
made  simultaneously  with  the  collection  of  the  organisms. 
By  this  means  one  element  in  the  behavior,  that  namely  of 
the  up-and-down  journeys  in  the  sea,  long  known  to  be  per- 
formed by  many  oceanic  species,  has  been  studied  with  a  fair 
degree  of  quantitative  accuracy  as  to  the  extent  of  travel, 
time  required  for  each  journey,  and  environmental  influence. 
A  considerable  series  of  species  have  been  compared  on  this 
basis. 


Organic  Connection  Between  Physical  and  Psf/chical     281 

Two  quotations  are  all  that  space  will  permit  us  to  give 
for  showing  what  these  investigations  are  bringing  out  on  tiie 
subject  of  specificity  in  behavior.  The  animals  referred  to 
in  the  first  quotation  constitute  a  group  of  small,  arrow- 
shaped  worms  known  as  the  Chaetognatha — bristle- jaws. 
"Each  species  occurring  in  the  San  Diego  region  has  its  own 
definite  and  specific  manner  of  vertical  distribution,  just  as 
truly  as  it  has  its  own  morphological  cliaracteristics."  -^ 

Similar  results  have  been  obtained  by  the  same  methods 
applied  to  a  very  different  group  of  animals,  minute  crus- 
taceans of  the  ubiquitous  order  Copepoda.  The  investiga- 
tions on  this  group  have  been  made  by  Dr.  C.  O.  Esterly, 
and  the  results  are  specially  interesting  in  this  case,  because 
Doctor  Esterly  has  applied  laboratory  experimentation,  to 
some  extent,  to  the  same  animals,  and  has  found  a  good  de- 
gree of  concordance  in  the  results  of  the  statistical  and  the 
experimental  investigations.  *'A  heterogeneous  assortment  of 
forms  may  be  obtained  in  the  same  collection  but  each  has  its 
own  way  of  reacting  toward  the  elements  of  its  environ- 
ment." 23 

It  is  the  indubitable  trend  in  one  direction  of  the  vast 
evidence  from  these  three  quite  different  classes  of  research 
on  animal  behavior  that  to  my  mind  justifies  such  a  con- 
ception of  specificity  of  psychical  and  reactive  animal  life 
as  that  formulated  above.  Something  of  the  probable  mean- 
ing of  this  specificity  we  shall  see  in  the  next  chapter. 

REFERENCE  INDEX 

1.  Loeb   ('02)    189  13.  Huxley,  J.  S 15G 

2.  Morgan,  C.  L 10  U.  Darwin,   C I,  353 

3.  Royce     328  15.  Kellogg,  V.   I litS 

4.  Royce     329  16.  Nature  Lovers'  Library  II,  218 

S.Holmes    ('11)     92  17.  Nature     Lovers'     Libnirv 

G.Holmes    ('11)    93                   III    100 

7.  Peckham,  G.  W.  and  E.  S.  53  IB.  Seton   I,  3«)0 

S.Holmes    ('11)    21  19.  Groos     15 

9.  Jennings    ('OG)    49  20.  Wheeler    ('10)     518 

10.  Holmes    ('11)     95        21.   Hitter   ('IS)    119 

1 1 .  Wallace,   A.    R 555       22.  Michael    4G 

12.  Huxley,  J.  S 151        23.  Esterly     11 


Chapter  XXIV 

SKETCH    OF    AN    ORGANISMAL    THEORY    OF 

CONSCIOUSNESS 

Remarks  on  the  Hypothetical  Character  of  This  Chapter 

HYPOTHESIS  and  theory  will  dominate  in  the  task 
upon  which  we  now  enter  and  in  this  respect  the 
present  chapter  will  differ  sharply  from  the  preceding  chap- 
ters. Fact,  description,  classification,  and  restrained  gen- 
eralization have  been  the  leading  motives  up  to  this  point. 
One  main  and  several  subsidiary  hypotheses  will  be  central 
in  the  discussion.  Into  the  presence  of  these  will  be  sum- 
moned many  of  the  facts  and  generalizations  previously  set 
forth.  The  purpose  in  this  summoning  will  be  on  the  one 
hand  to  test  the  hypotheses  by  the  facts  and  generalizations 
and  on  the  other  hand  to  see  how  the  facts  will  look  in  the 
light  of  the  hypotheses. 

This  announcement  of  the  hypothetical  and  theoretical 
character  of  the  task  now  before  us,  will  give  us  two  advan- 
tages:  It  will  justify  a  dogmatic  form  of  expression  at  times 
which  we  should  not  otherwise  feel  privileged  to  use;  and 
will  justify  a  brevity  of  treatment  which  would  not  be  pos- 
sible were  we  aiming  at  thorough  generalization  and  demon- 
stration. Hence  the  justification  of  undertaking  to  deal 
with  so  vast  and  vital  a  subject  in  the  limits  of  a  sketch. 

The  Natural  History  Method  and  the  Stiidy  of  One's  Self 

Insistent  as  I  have  been  on  the  importance  of  the  natural 
history  wa}^   of  approaching  the   phenomena   of  the  living 

282 


Sl-etch  of  an  Organismal  Theory  of  Cori.u'wnsnfss     283 

world,  in  entering  upon  tlie  present  discussion  I  must  em- 
phasize this  more  than  ever  and  nuist  call  attention  to  tlie 
particular  character  of  this  importance  in  our  present  un- 
dertaking-. 

The  natural  history  method  of  viewing  organic  hiings  is 
2)er  se  the  comprehensive  method,  one  of  its  best  mottos 
being,  as  we  have  repeatedly  seen,  "neglect  nothing."  Tliat 
knowledge  of  organisms  separates  itself  sharply  into  de- 
partments is  no  deterrent  to  the  naturalist  against  utihzing 
any  knowledge  he  may  come  upon  that  will  contribute  to  his 
main  aim — that  of  understanding  organisms.  \Yho  or  what 
shall  restrain  me  from  observinn^  and  carefully  thinkine: 
about  any  fact  of  my  own  being  which  promises  to  help  me 
on  my  road  to  such  understanding.'^  The  foremost  zoolo- 
gists, of  modern  times  especially,  have  amply  recognized  and 
freely  used  this  principle  so  far  as  all  physical  and  some  of 
the  lower  psychical  attributes  are  concerned.  But  when  it 
comes  to  man's  higher  psychical  attributes,  zoologists  have 
usually  said,  sometimes  expressly,  sometimes  tacitly,  that 
these  belong  to  a  wholly  different  realm,  a  realm  with  which 
we  have  little  or  nothing  to  do.  And  their  position  of 
"hands  off"  as  touching  man's  higher  psychic  life,  has  re- 
ceived the  readier,  fuller  sanction  in  that  it  has  accorded 
well  with  the  prevalent  views  and  practices  of  those  students, 
anthropologists,  economists,  sociologists,  and  ethicists  who 
have  made  these  higher  reaches  of  human  life  their  special 
fields  of  inquir}'.  But  the  course  of  nature  can  not  be  per- 
manently thwarted.  Such  an  attempt  to  wrench  human  life 
asunder  is  bound  to  fail  finall3\  In  tlie  several  sulxlivisions 
of  biology,  normal  advance  has  tended  to  stay  the  wrencliiiig 
process,  comparative  psychology  being  notable  in  this  ten- 
dency. 

The  opposition  to  such  organic  disunion  consistently 
maintained  throughout  this  book  reaches  its  culmination  in 
these  chapters  on  psychic  integration.     In  what  follows  we 


284  The  Unity  of  the  Organism 

shall  pass  more  freely  than  ever  from  one  phase  or  aspect 
to  another,  over  the  entire  gamut  of  psychic  life  both  in  the 
individual  and  in  the  animal  kingdom.  If  facts  of  my  own 
subjective  life  will  serve  my  purpose,  I  shall  be  as  free  to 
requisition  them  as  to  requisition  facts  of  any  phase  or  as- 
pect of  my  objective  life.  If  the  ethical  or  esthetic  or  social 
attributes  of  the  human  animal  will  best  illuminate  a  point, 
these  shall  be  brought  in  with  as  little  misgiving  as  will  be 
anatomical  or  embryological  or  physiological  or  instinctive 
attributes. 

So  great  store  do  I  lay  on  this  catholicity  of  attitude 
toward  psychic  life,  that  I  shall  show  by  a  single  instance 
that  at  least  a  few  other  present-day  zoologists  have  some- 
what similar  feelings  about  the  zoological  character  of  psy- 
chical phenomena.  Referring  to  the  controversies  which 
have  inevitably  arisen  over  the  problem  of  instinct,  W.  M. 
Wheeler  says  that  such  controversy  "is  pardonable,  at  least 
to  some  extent,  since  the  subject  itself  presents  no  less  than 
four  aspects,  according  as  it  is  studied  from  the  ethological, 
physiological,  psychological  or  metaphysical  points  of  view." 
"From  the  first  two  of  these,"  the  author  continues,  "in- 
stinct is  open  to  objective  biological  study  in  the  form  of 
the  'instinct  actions.'  These  may  be  studied  by  the  physiol- 
ogist merely  as  a  regularly  coordinated  series  of  movements 
depending  on  changes  in  the  tissues  and  organs,  and  by  the 
ethologist  to  the  extent  that  they  tend  to  bring  the  organism 
into  effective  relationship  with  its  living  and  inorganic  en- 
vironment. But  that  these  movements  have  a  deeper  origin 
in  psychological  changes  may  be  inferred  on  the  basis  of 
analogy  from  our  own  subjective  experience  which  shows  us 
our  instincts  arising  as  impulses  and  cravings,  the  so-called 
'instinct-feelings';  and  these  in  turn  3aeld  abundant  material 
for  metaphysical  and  ethical  speculation."  ^  From  the 
context  of  these  sentences  we  may  infer  that  Wheeler  rec- 
ognizes  that   the   four  aspects   mentioned   under  which  the 


Sketch  of  an  Organismal  Theory  of  Consciousmss     285 

subject  of  instincts  presents  itself,  represent  the  same  num- 
ber of  valid  departments  of  man's  mental  life.  The  point 
I  wish  to  make  is  that  although  a  zoologist  may  recognize 
without  cavil  that  speculation  on  psychological,  ctliical, 
and  metaphysical  problems  which  arise  in  connection  witli 
instincts,  are  legitimate  activities  of  man,  and  might  prop- 
erly deny  that  it  is  incumbent  upon  him  to  do  much  specu- 
lating of  this  sort,  yet  it  would  be  incumbent  on  him  to  take 
due  cognizance  of  these  speculative  attributes  of  the  human 
animal.  A  truly  scientific  zoology  can  not  justify  itself  in 
issuing  a  manifesto  to  the  effect  that  certain  attributes  pre- 
sented by  some  animals  do  not  fall  within  its  province.  It 
may  more  or  less  constantly  neglect  or  refuse  on  practical 
grounds,  to  deal  with  certain  attributes ;  but  that  is  a  very 
different  matter  from  a  formal  declaration  such  as  many 
present-day  zoologists  make,  that  with  these  attributes 
zoology  has  nothing  to  do.  Such  a  declaration  is  self-stunt- 
ing, if  not  self-stultifying,  in  that  it  is  a  virtual  self-inhi- 
bition by  zoology  of  its  own  growth. 

These  reflections  may  be  terminated  by  defining  the  mo- 
tives and  the  mental  attitude  with  which  I  approach  the 
great  problem  of  consciousness.  I  come  to  it  not  as  a  meta- 
physician, not  as  a  psychologist,  not  as  a  physiologist,  not 
even  as  an  anthropologist,  but  as  an  anthrojjological  zoolo- 
gist ;  as  a  zoologist  who  in  course  of  his  regular  professional 
work  takes  up  the  animal  group  of  which  he  himself  is  a  mem- 
ber, chancing  as  he  does  to  possess  among  other  attributes 
that  of  knowing  his  own  life  directly,  that  is,  through  sub- 
jective or  self-conscious  experience,  as  well  as  indirect  I  if 
through  objective  experience. 

Approaching  the  problem  of  consciousness  in  such  an  at- 
titude and  for  such  a  motive,  it  is  impossible  to  view  it  other- 
wise than  as  one  aspect  of  the  larger  problem  of  life  gener- 
ally. For  while  the  psychologically  and  metaphysically  im- 
portant   question    of    whether    consciousness    is    coextensive 


286  The  Unity  of  the  Organism 

with  life  need  not  be  raised  by  the  naturalist,  the  indubitable 
fact  that  at  least  a  large  sector  of  life  is  conscious ;  in  other 
words,  the  fact  that  consciousness  is  a  part  of  life,  he  can 
not  ignore  if  he  is  to  deal  with  consciousness  at  all.  For  the 
naturalist,  then,  no  hypothesis  or  theory  of  consciousness 
can  be  satisfactory  which  is  not  clearly  and  expressly  em- 
bedded in  and  an  essential  part  of  an  hypothesis  or  theory 
of  life  generally.  Our  central  hypothesis,  drafted  in  ac- 
cordance With,  these  principles,  may  now  be  given. 

Formulation  of  the  Central  Hypothesis 

All  the  manifestations  which  in  the  aggregate  we  call 
Life,  from  those  presented  hy  the  simplest  plants  to  those 
of  a  consciously  psychical  nature  presented  hy  man  and 
numerous  other  animals,  result  from  tJie  chemical  reaction 
between  the  organism  and  the  respiratory  gases  they  take, 
oxygen  being  almost  certainly  the  effective  gas  for  nearly 
all  animals.  An  essential  implication  of  this  proposition 
is  that  every  living  individual  organism  has  the  value, 
chemically  speaking,  of  an  elementary  chemical  substance. 

Let  us  be  promptly  explicit  in  recognizing  the  character 
of  the  two  propositions  contained  in  this  hypothesis.  They 
are  manifestly  chemical  in  large  part,  and  a  complete  demon- 
stration of  their  truth  is  impossible  without  the  aid  of  chem- 
ical research  focus sed  directly  upon  them.  But  though 
clearly  chemical,  equally  clearly  they  go  beyond— far  beyond 
— present  chemical  knowledge.  To  speak  of  a  whole  organ- 
ism as  equivalent  to  a  chemical  element  seems  at  first  sight 
not  only  unwarranted  by  positive  chemical  knowledge,  but 
opposed  by  such  knowledge.  Furthennore,  the  term  "re- 
action" as  used  in  the  first  proposition  undoubtedly  seems 
quite  foreign  to  the  technical  meaning  which  chemistry  has 
attached  to  the  word.  Indeed  so  remote  to  say  the  least, 
are  these  fundamental  propositions  of  the  hypothesis  from 


Sketch  of  an  Organismal  Theory  of  ConsciouJiuess     287 

definite  chemical  knowledge,  that  if  they  are  entitled  to  rank 
as  constituting  a  legitimate  scientific  hypothesis,  this  must 
be  on  grounds  other  than  those  of  present-day  technical 
chemistry  quite  as  much  as  on  those  of  such  chemistry.  In 
attempting,  consequently,  to  establish  the  propositions  on  a 
tiTie  and  useful  hypothetical  basis,  it  will  be  permissible  to 
notice  these  other  grounds  first. 

Preliminary  Justification   of   the   Hypothesis   as  Such 

The  proposition  that  each  living  individual  has  the  chem- 
ical value  of  an  elementary  substance,  will  receive  attention 
first,  and  the  initial  step  will  be  to  inquire  what,  in  general, 
the  criterion  is  of  an  elementary  chemical  substance.  Here, 
for  instance,  is  a  lump  of  phosphorus.  In  virtue  of  what  is 
it  declared  to  be  such  a  substance  .^  Not  primarily,  let  us 
specially  notice,  because  the  phosphorus  is  simple,  that  is 
to  say,  is  an  element  in  the  sense  of  not  being  reducible  to 
still  simpler  substances.  Rather  the  basal  criterion  of  its 
being  a  chemical  substance  is  that  upon  its  being  brouglit 
into  contact  under  certain  conditions  with  certain  other 
chemical  substances,  oxygen  for  instance,  there  is  produced 
a  third  substance  having  very  different  attributes  from  either 
of  the  original  substances.  Transformation  of  substances 
chiefly  through  interaction  upon  one  another  is  the  founda- 
tion fact  which  has  brought  it  to  pass  that  substances  are 
described  as  chemical.  That  is  the  fact  upon  which  the 
science  of  chemistry  primarily  rests.  Facts  and  problems 
of  simplicity  and  complexity,  relative  and  absolute,  are 
later  and  secondary.  The  task  of  chemistry  "consists  in  the 
investigation  of  substances  and  those  of  their  processes  by 
which  the  physical  attributes  of  the  substances  undergo 
permanent   dianges."      {Hanchcoricrhiicli    drr  Xafur-cissm- 

schaft.) 

Every  adequate  definition  of  chemistry  and  chemical  sub- 


288  The  Unity  of  the  Organism 

stance  and  chemical  action  contains  the  idea  of  transforma- 
tion in  one  form  or  another.  Clearness  on  this  point  is  in- 
dispensable to  our  purpose.  Chemistry  is  too  often  defined, 
even  in  elementary  text  books  and  in  dictionaries,  as  though 
the  "composition  of  matter"  were  its  initial  and  most  es- 
sential function.  But  this  conception  is  surely  contrary  to 
the  history  and  most  essential  nature  of  the  science.  There 
is,  it  seems,  entire  agreement  among  competent  writers  that 
scientific  chemistry  is  a  direct  descendant  of  Alchemy,  and 
a  very  imperfect  knowledge  of  the  history  of  Alchemy  re- 
veals the  fact  that  the  every-where  present,  normal  trans- 
formations in  nature,  particularly  in  inorganic  nature,  were 
the  foundation  phenomena  of  this  old  art.  One  has  only  to 
recall  the  place  held  by  the  idea  of  the  transmutation  of 
metals,  this  idea  having  usually  the  practical  aim  of  chang- 
ing the  "base  metals"  into  "noble  metals,"  The  "phil- 
osopher's stone"  and  the  "great  elixir"  were  magical  some- 
tliings  by  which  the  transmutations  could  be  accomplished. 

Greatly  significant  from  our  standpoint  is  the  fact  that 
one  of  the  objectives  of  Robert  Boyle  (middle  of  the  seven- 
teenth century),  who,  perhaps  as  much  as  any  one  man,  is 
entitled  to  be  called  the  father  of  experimental  chemistry, 
was  to  rectify  the  false  and  mystical  notions  prevalent  in  his 
time  about  "Elements,"  "Principles,"  "Essences,"  etc.  "Tell 
me  what  you  mean  by  your  Principles  and  your  Elements," 
Boyle  demanded,  "then  I  can  discuss  them  with  you  as  work- 
ing instruments  for  advancing  knowledge."  ^ 

What  is  "behind"  the  transformations — forces,  elements, 
principles,  essences,  spirits  or  what  not — is  indeed  an  impor- 
tant and,  properly  asked,  a  legitimate  question.  But — and 
here  is  the  most  vital  fact  of  all — it  is  a  question  which  can 
not  be  raised  even,  until  after  the  transformations  have  been 
observed,  nor  can  an  answer  of  objective  value  be  given  un- 
less the  whole  round  of  observed  phenomena,  the  substances 
previous  to  transformation,  the  transformatory  processes. 


Sketch  of  an  Organhmal  Theory  of  Consciousness     289 

and  the  new  substances,  be  accepted  at  their  face  value,  that 
is  to  say,  at  a  value  which  is  as  near  to  ultimate  truth  as  any 
truth  whatever,  connected  with  the  phenomena. 

The  elemental  constitution  of  bodies  is  an  inference,  al- 
ways and  solely,  drawn  from  their  observed  corporeal  attri- 
butes. And  chemistry  is  the  science  which  assumes  the  task 
of  drawing,  elaborating,  and  systematizing  these  inferences 
on  tlic  basis  of  the  transformation  of  the  attributes.  Tlu- 
meaning  of  the  statement  that  chemistry  is  one  of  the  natural 
sciences  is  that  chemistry  is  the  science  which  uses  its  natural 
history  observations  to  penetrate  still  more  deeply  into  the 
constitution  of  bodies.  Natura  a  nutwra  vincitury  nature  is 
surrounded  by,  is  contained  in  nature,  is  as  fundamental  a 
truth  for  chemistry  as  for  any  other  natural  science.  A 
living  being  is  as  much  a  natural  body  as  is  a  piece  of  phos- 
phoi*us,  and  its  obvious  attributes,  its  outer-layer  attributes, 
are  as  essential  to  its  nature  as  are  its  inner,  its  hidden 
attributes.  So  any  genuinely  transformatory  changes,  and 
genuinely  new  products  arising  through  the  reaction  between 
the  living  body  and  some  other  body  is  so  far  chemical  in 
nature,  and  the  reacting  bodies  are  so  far  chemical. 

A  long  step  toward  justifying  the  proposition  that  each 
individual  living  organism  has  the  value,  chemically,  of  an 
elementary  substance,  will  be  taken  if  it  can  be  shown  that 
any  qualitatively  new  product  whatever  results  from  the 
interaction  between  the  organism  acting  as  a  unit,  as  one, 
as  an  element,  and  some  other  element.  Having  regard  to 
the  entire  world  of  living  beings,  the  chances  for  finding  new 
products  which  may  have  arisen  in  this  manner  are  ]>rac- 
tically  if  not  theoretically  infinite.  IManifestly,  tlien,  only  a 
very  small  sector  of  the  entire  range  of  such  j)()ssible  produc- 
tions can  be  searched.  It  must,  consequently,  Ik'  our  aim,  as 
always  in  handling  inductive  natural  history  evidence,  to 
choose  for  examination  evidence  which  shall  be  most  clear- 
cut,  most  illustrative,  and  most  convincing. 


290  The  Unity  of  the  Organism 

The  sector  of  organic  phenomena  best  capable  of  yielding 
such  evidence  is,  I  believe,  exactly  this  of  psychic  life.  And 
within  the  great  range  of  this  life,  the  higher  conscious  life 
of  man  is  most  replete  with  the  evidence  we  seek.  Again 
within  the  range  of  man's  higher  life,  each  individual's  own 
private  life,  even  his  subjective  life,  his  consciousness,  is  the 
evidence  most  certain  and  convincing.  Translating  this  last 
statement  into  familiar  language,  one  sees  that  it  is  only 
another  way — the  scientific  way — of  affirming  the  truth,  that 
the  greatest  of  all  certainties  of  which  man  is  capable  is  that 
of  his  own  existence.  I  am  saying,  virtually,  that  when  we 
analyze,  after  the  manner  of  objective  science,  this  old  fa- 
miliar affirmation  about  certainty,  and  carry  the  analysis 
as  far  as  we  are  at  present  able  to,  we  find  that  the  sense,  or 
better,  the  feeling  of  certainty  of  self-existence  and  self- 
identity  is  in  last  analysis  one  of  the  effects  of  a  transforma- 
tory  interaction  between  ourselves  and  some  substance  (oxy- 
gen?) in  our  breath,  as  stated  in  the  first  of  our  two  propo- 
sitions. 

That  proposition  seems  then  to  be  hardly  more  than  a 
recognition  that  psychic  phenomena  containing  at  least  the 
germ  of  consciousness  is  a  kind  of  chemical  product  which 
has  not  heretofore  been  clearly  recognized  as  such,  the  lack 
of  recognition  being  due  to  the  strangeness  of  the  product  as 
compared  with  any  chemical  products  with  which  experimen- 
tal chemistry  has  hitherto  occupied  itself.  But  looked  at  in 
a  really  broad  and  deep  way,  is  it  any  more  difficult  for  me 
to  interpret  a  state  of  consciousness  in  myself  to  be  a  result 
af  chemical  action  between  me  and  the  air  (oxygen .f*)  I 
breathe,  than  for  me  to  interpret  the  dim  greenish-white 
luminosity  of  a  piece  of  phosphorus  to  be  a  result  of  the 
chemical  action  between  the  phosphorus  and  the  air  essential 
to  the  glowing?  From  a  purely  chemical  standpoint  I  do 
not  believe  we  have  any  ground  for  holding  that  some  prod- 
ucts of  chemical  reaction  are  more  comprehensible  or  less 


SletcJi  of  an  Organismol  Thcori/  of  Consciousiuss     1^291 

comprehensible  tlian  are  otliers. 

Chemically  viewed  tlie  problem  now  on  our  hands  is  en- 
tirely one  of  fact — fact  as  determined  by  observation  alone, 
and  by  observation  with  the  aid  of  experimentation.  If  it 
can  be  shown  that  each  individual  conscious  beinr^  really  does 
behave  like  a  chemical  substance  in  the  process  of  reacting; 
and  if  the  result  of  such  reaction  can  be  shown  to  have  even 
one  of  the  essential  marks  of  a  chemical  product,  both  propo- 
sitions of  my  two-parted  hypothesis  are  warrantable  and  the 
hypothesis  becomes  genuinely  scientific — a  genuine  "working 
hypothesis" — one,  that  is,  for  bio-chemistry  to  take  seri- 
ously. 

More  Systematic  Justification  of  the  Hypotliesis 

That  the  propositions  are  demonstrable  to  the  extent  of 
the  demand  just  indicated  is  my  contention.  Tliis  conten- 
tion I  will  now  try  to  make  good  and  will  begin  with  a  few 
remarks  on  a  question  concerning  the  hypothesis  which 
ought  to  arise  instinctively  in  the  mind  of  every  one.  That 
question  is :  Does  such  a  conception  of  psychic  life  and  con- 
sciousness as  that  contained  in  our  hypothesis  imply  any  real 
infringement  upon  or  derogation  from  me,  in  the  deepest 
sense  a  real  entity  properly  designated  by  the  terms  -person 
and  personality? 

On  saying  that  this  query  ought  to  arise  insfinctivrli/,  I 
do  not  mean  ought  in  the  ethical  sense,  but  in  the  organismal 
sense.  That  is,  in  a  sense  which  imj)lies  that  the  very  nature 
of  the  conscious  organism  is  that  it  is  not  only  self-existent 
in  a  measure  like  every  natural  object,  but  that  it  is  self-iden- 
tifiable, and  within  certain  bounds,  self-determinative  of  its 
own  acts.  Now  recognizing  it  to  Ix?  thus  by  its  "very  na- 
ture" is  only  another  way  of  recognizing  that  it  is  so  in  its 
instincts  as  well  as  in  its  physical  organization.  But  since 
instinct  is  more  fundamental,  more  deep-rooted  in  the  or- 


29^  The  U7iity  of  the  Organism 

ganism  than  is  intellect,  as  phylogenic  and  ontogenic  psy- 
chology make  clear,  if  a  pronouncement  implying  a  de- 
rogation from  the  reality  and  natural  prerogatives  of  the 
individual  be  issued  from  the  intellect,  a  response  of  protest 
and  antagonism  would  be  expected  from  instinct.  This  would 
be  expected  as  an  ordinary  organic  impulse  to  self-defense 
and  self-preservation. 

The  Nature  of  ''Outer''  or  Objective  and  "Inner'*  or 

Subjective 

What  we  have  to  do  consequently  is  to  scrutinize  the  con- 
scious individual  in  order  to  see  if  it  presents  any  uniqueness 
of  attributes  and  of  transformatory  power  in  reacting  with 
other  bodies  that  is  on  a  par  with  the  uniqueness  of  an  ordi- 
nary chemical  substance  in  the  same  respects.  Now  it  is, 
as  suggested  some  pages  back,  exactly  in  the  conscious,  the 
subjective  life,  that  such  uniqueness  is  most  easily  demon- 
strable. There  are  several  ways  in  which  the  conscious  indi- 
vidual manifests  this  uniqueness.  A  particularly  convincing 
way,  I  think,  is  in  the  relation  between  w^hat  are  commonly 
known  as  the  objective,  or  "outer,"  and  the  subjective  or 
"inner"  sides  of  mental  life.  This,  consequently,  will  be  the 
approach  to  the  subject  chosen  by  us  and  we  will  enter  upon 
it  by  returning  to  Royce,  first  to  his  "OutHnes  of  Psychol- 
ogy," then  a  little  later  to  some  of  his  specifically  philosophi- 
cal writings. 

In  the  first  chapter  of  the  Outlines,  devoted  to  initial  defi- 
nitions and  explanations,  Royce  states,  simply  and  clearly, 
a  distinction  "between  our  physical  and  mental  life,"  which 
elsewhere  he  has  worked  out  with  great  elaboration.  Thus: 
"Physical  facts  are  usually  conceived  as  'public  property,' 
patent  to  all  properly  equipped  observers.  All  such  observ- 
ers, according  to  our  customary  view,  see  the  sarne  physical 
facts.    But  psychical  facts  are  essentially  'private  property,' 


Sketch  of  an  Orgamsmal  Thcor//  of  Conscion.sfuss     ^298 

existent  for  one  alone.  This  constitutes  the  very  conception 
of  the  difference  between  'inner'  psycliical  or  mental,  and 
physical  or  'outer'  facts.""* 

Ever-present,  and  obvious  as  is  the  comparison  here  made, 
it  nevertheless  is  of  so  great  importance  that  we  must  stop 
and  reflect  upon  it,  for  we  shall  surely  fail  to  grasp  the  full 
measure  of  what  is  to  follow  if  we  are  lukewarm  toward  one 
of  the  elements  of  it.  The  element  I  refer  to  is  the  unique- 
ness, the  essentially  personal  character  of  inner  as  contrast- 
ed with  outer  facts.  Every  normal  person  is  ready  enough 
to  insist  that  his  thoughts,  his  feelings,  his  emotions  and  all 
the  rest  of  his  higher  psychical  experiences  are  his  and  his 
alone.  The  tremendous  reality  and  force  of  the  rights  of 
"private  opinion,"  of  "personal  conscience"  and  so  forth, 
among  civilized  men,  hardly  need  to  be  expatiated  on. 

The  character  of  the  uniqueness  of  these  experiences,  how- 
ever, concerns  practical  living  less  vitally,  so  we  give  it  less 
attention.  The  whole  vast  range  of  my  mental  life,  from 
the  lowest,  simplest,  vaguest  sensations  to  the  highest,  most 
bewildering  complex  emotions,  passions,  imaginings  and 
thoughts,  are  my  own,  absolutely,  so  far  as  other  persons 
are  concerned.  I  cannot  share  them  to  the  least  extent  with 
another  person.  Of  course  I  can  let  others,  especially  my 
most  intimate  associates,  m}^  dearest  friends,  know  a  good 
deal  about  these  experiences  of  mine.  But  after  all,  gladly 
as  I  would  share  many  of  them  with  these  friends,  it  is  utter- 
ly impossible  for  me  to  do  so.  Mi/  experiences  must  remain 
wholly  outside  of  their  consciousness.  No  two  persons  can 
have  the  same  experience  an}^  more  than  they  can  have  the 
same  hands  or  stomachs.  Nor  is  this  all.  If  mental  life  is 
subject  to  the  general  biological  laws  of  variation  into  whieh 
we  have  latterly  gained  much  insight,  I  am  obliged  to  sup- 
pose that  these  experiences  of  mine,  the  whole  retiiuie  of  sen- 
sations, feelings,  emotions  and  thoughts,  differ  somewhat 
from  the  corresponding  experiences  of  other  persons.     And 


^94  The  Unity  of  the  Organism 

all  observation  confirms  this  supposition — much  of  it  strong- 
ly. Inferential  evidence  could  hardly  be  stronger  than  that 
my  particular  emotional  response  to  opera  singing,  for 
example,  is  quite  different  from  that  of  many  other  persons. 
Obviously  we  are  here  skirting  the  edge  of  what  modern 
realism  in  formal  philosophy  calls  pluralism,  and  deals  with 
in  part  as  the  question  of  whether  percepts  are  strictly  indi- 
vidual and  personal.  No  pliilosopher  with  whose  views  I 
have  become  acquainted,  has  discussed  this  question  so  fully, 
and  in  my  opinion,  so  illuminatingly  as  Sellars.  The  follow- 
ing sentences  taken  from  his  chapter,  The  Advance  of  the 
Personal,  show  clearly,  it  seems  to  me,  that  the  conclusions 
he  has  reached,  working  from  the  purely  philosophical  side, 
are  essentially  the  same  as  those  arrived  at  by  me,  advancing 
from  the  biological  side:  "What  may  be  called  the  sensory 
content  of  our  percepts  is  important, — I  do  not  wish  to 
be  understood  to  belittle  it, — but  so  are  the  meanings  which 
arise  in  conection  with  our  bodily  activities  and  motor  ad- 
justments to  stimuli.  Here  again,  we  are  face  to  face  with 
individual  factors  in  perception  which  even  the  idealist  must 
recognise  and  somehow  explain.  Evidently,  perception  is  not 
a  mere  passive  presentation,  but  a  construction  whose  gene- 
tic elements  can  be  partially  traced.  Finally,  let  us  call  to 
mind  that  percepts  are  continuous  with  feelings  and  with  the 
so-called  organic  sensations.  .  .  .  Once  vaguely  objective, 
feeling  is  now  considered  subjective  or  personal."^"^  Many 
other  sentences  and  paragraphs  of  like  purport  could  be 
quoted  from  this  author.  I  have  selected  this  for  the  two- 
fold reason  that  it  indicates  the  measure  of  my  agreement 
with  his  view  as  to  the  personal  character  of  percepts  and 
the  rest  of  conscious  life;  and  at  the  same  time  indicates 
wherein  I  shall  have  to  out-do  him  in  the  matter  of  validat- 
ing the  individual.  A  part  of  our  task,  to  be  reached  a  little 
later,  will  be  to  show  that  although  feeling  and  all  the  rest 
of   psychic   life   is   indeed   subjectively   personal,   it   is    also 


Sketch  of  an  Orgamsvinl  Theory  of  Conscious luss     Ji95 

objectively  personal.  In  otlicr  words,  it  will  })l'  my  task  to 
remove,  or  at  least  to  show  the  way  to  remove,  the  vagueness 
which  Sellars  asserts,  rightly,  has  hitherto  cloudid  this 
side  of  personality.  To  do  this  thing  is,  indeed,  one  of  my 
most  important  chances  to  contribute  to  a  "better  philosoj)hy 
of  life." 

But  since  our  psychical  life,  especially  our  conscious  life, 
is  a  vast — incalculably  vast — complex  of  experiences,  of 
"contents,"  sounds,  sights,  memories,  feelings,  ideas,  many 
of  which  are  set  off  very  sharply  from  the  rest,  are  clearly 
characterizable,  and  are  wonderfully  2:)ersistent ;  and  since 
innumerable  of  these  are  coming  along  all  the  while  which 
have  much  of  genuine  newness  about  them ;  and  since  further, 
these  contents  of  consciousness  are  intertwined  with  and  are 
determinative  of  a  vast  complex  of  other  contents  called  voli- 
tions which  in  turn  are  linked  up  with  and  are  more  or  less 
directive  of  bodily  activities  of  many  kinds,  some  purely  re- 
flex and  some  instinctive,  it  seems  impossible  to  escape  recog- 
nizing, even  if  one  wanted  to,  that  if  the  verb  "to  create"  has 
any  definite  meaning  at  all  the  normal,  self-conscious  animal 
organism  is  about  the  most  creative  thing  we  know  or  can 
conceive.  Indeed  it  is  altogether  likely  that  the  very  notion  . 
of  creation,  whether  natural  or  supernatural,  came  initially 
from  the  creative  activity  and  the  impulse  to  such  activity, 
of  man  himself. 

We  may  justly  say,  I  think,  that  we  know  all  creativeness, 
chemical  creativeness  with  the  rest,  through  being  in  our  own 
deepest  natures  creative,  that  is,  transformative  and  trans- 
formative in  the  way  which  we  call  chemical.  We  learn 
about  the  processes  of  life  and  call  some  of  the  most  essen- 
tial of  them  chemicjal  just  by  pciiorming  those  ])rocesses  as 
some  of  our  most  essential  attributes.  A  portion  of  the  ]iro- 
cess  which  goes  on  within  us,  together  with  the  corresponding 
product,  constitutes  what  we  call  the  science  of  bio-chemis- 
try.    This  means  that  according  to  our  hypothesis  "objcc- 


296  The  Unity  of  the  Organism 

the"  and  "subjective,"  or  "outer"  and  "inner"  as  applied  to 
life,  are  something  quite  different  from  what  they  have  been 
either  in  traditional  philosophies,  or  in  most,  at  least,  of 
recent  psychology.  "When  we  speak,"  Royce  writes,  "of 
our  physiological  processes  as  internal,  the  word  'internal,' 
although  it  here  generally  implies  'hidden,  in  whole  or  in  part, 
from  actual  outer  observation',  does  not  imply  'directly  felt 
by  us  ourselves.'  "  ''  My  hypothesis  implies  a  denial  of  the 
correctness  of  this  statement.  I  say  that  in  the  sum  total 
of  the  "contents  of  consciousness,"  a  nether  segment,  as  one 
might  call  it,  of  physiological  processes  is  "directly  felt  by 
ourselves."  There  is  no  content  of  consciousness  which  does 
not  contain  an  element  that  is  internal  or  subjective  in  what- 
ever sense  any  other  content  of  consciousness  is  internal  ov 
subjective.  And  per  contra,  there  is  no  content  of  conscious- 
ness which  is  not  objective  to  some  extent,  in  whatever  sense 
any  other  content  of  consciousness  is  objective.  The  mind, 
according  to  this  conception,  is  not  something  which  uses 
the  brain  or  any  other  part  of  the  organism  merely  as  a  tool 
with  which  to  make  thoughts  and  other  contents  of  con- 
sciousness. Nor  on  the  other  hand  is  consciousness  of  the 
nature  of  a  secretion,  the  gland  for  which  is  the  brain,  though 
unquestionably  the  brain  has  an  essential  part  in  the  pro- 
duction of  thought  and  the  higher  contents  of  consciousness. 
Among  the  consequences  of  the  reaction  between  the  or- 
ganism and  the  air  we  breathe  are  consciousness  with  its 
marvellously  rich  and  varied  contents. 

But  at  this  point  I  must  specially  request  the  reader  to 
notice  that  I  am  not  pretending  to  describe  and  explain  all 
the  contents  of  consciousness.  In  other  words  it  is  not  a 
theory  of  knowledge^  but  a  theory  of  consciousness  that  I 
am  sketching;  and  knowledge  in  the  strict  sense,  and  con- 
sciousness are  very  different.  They  differ,  according  to  my 
understanding,  much  as  the  fully  developed,  physical  organ- 
ism  differs    from   the   living   substance,   or   protoplasm,    of 


Sketch  of  an  Organismnl  Theorif  of  Con.sciousmss     297 

which  the  organism  is  composed.  Consequently  I  am  not 
even  concerned  primarily  with  sensation  in  so  far  as  this  im- 
plies sense  organs  or  even  nerves  and  nerve  terminals  of  the 
simplest  kind.  Rather  1  am  dealing  witii  the  stages  and  con- 
ditions antecedent  to  consciousness  and  in  which  it  is  latent, 
in  much  such  way  as  the  cytologist  when  he  studies  the  living 
substance  of  all  sorts  of  tissue-cells  is  not  dealing  with  organs 
and  the  organism  in  the  full  sense,  but  only  with  their  sub- 
strata. But  although  it  is  not  knowledge,  properly  speaking, 
either  in  its  conceptual  or  perceptual  aspect  that  I  am  dis- 
cussing, since  my  enterprise  does  take  me  across  the  border 
line  and  a  short  distance  into  the  realm  of  knowledge,  I  must, 
in  the  interest  of  historical  continuity  and  setting,  say  a 
little  more  than  I  have  said  about  the  general  nature  of 
knowledge. 

My  assertion  should  be  taken  literally  that  there  is  no 
content  of  consciousness  which  is  purely  either  subjective  or 
objective,  inner  or  outer,  conce})tive  or  perceptive,  ideational 
or  impressional,  or  whatever  form  of  expression  be  given  the 
antithesis  here  implied.  That  every  content  of  consciousness 
which  exists  or  can  be  conceived  has  an  essential  element  of 
both  members  of  the  antithesis  is  exactly  what  I  mean.  To 
illustrate,  even  the  axioms,  postulates,  or  whatever  else  may 
be  counted  as  most  ultimate  in  mathematics  contain  an  ele- 
ment of  the  outer,  or  objective,  as  well  as  of  the  inner,  or 
subjective.  These  mathematical  contents  of  consciousness  I 
single  out  to  illustrate  my  meaning  because  they  have  been 
clung  to  by  philosophers  and  scientists  more  tenaciously  than 
any  others  as  purely  subjective  or  mental.  And  further 
there  is  a  strategic  gain  in  this  reference  to  mathematics  in 
that  it  brings  into  the  open  the  fundamental  opposition  of 
my  hypothesis  to  one  main  root  of  Cartesian  i)l»ilosoj)liy  ; 
the  philosophy,  that  is,  from  which  the  modern  doctrine  of 
psycho-physical  parallelism  has  grown.  Our  thinking,  which 
Descartes  held  proves  our  existence,  really  proves  it  only  in 


^98  The  Unity  of  the  Organism 

so  far  as  it  shows  that  among  the  activities  essential  to  the 
human  organism  thinking  is  one.  In  other  words  the  "there- 
fore" in  "I  think,  therefore  I  am,"  is  true  only  because  *'I 
am,  therefore  I  think,"  the  reverse  proposition,  is  also  true 
and  includes  the  other  truth.  The  lesser  truth  is  true  be- 
cause it  is  an  essential  part  of  the  larger  truth,  much  in 
the  same  way  that  the  cells  of  a  multicellular  organism  are 
alive  because  they  are  essential  parts  of  the  organism. 

We  need  not  inquire  how,  from  this  serious  shortcoming  of 
Descartes'  description  of  psychic  life  Descartes  went  on  to 
the  conclusion  that  "there  is  nothing  really  existing  apart 
from  our  thought"  and  that  "neither  extension,  nor  figure, 
nor  local  motion,  nor  an^^thing  similar  that  can  be  attrib- 
uted to  body,  pertains  to  our  nature,  and  nothing  save 
thought  alone ;  and,  consequently,  that  the  notion  we  have  of 
our  mind  precedes  that  of  any  corporeal  thing,  and  is  more 
certain,  seeing  we  still  doubt  whether  there  is  any  body  in 
existence,  while  we  readily  perceive  that  we  think."^^  Nor 
need  we  concern  ourselves  with  the  voluminous  and  tedious 
reasonings  by  which  a  considerable  number  of  moderns,  fol- 
lowing Descartes's  lead,  have  convinced  themselves  that  they 
have  "reduced"  all  realit}'  or  at  least  all  reality  that  really 
amounts  to  anything,  to  quantity.  Enough  now  to  remark 
that  every  modern  biologist  who  really  accepts  the  basal  data 
of  his  science,  must  agree  that  "Psycho-physical  paralellism 
.  .  .  stands  to-day  as  the  scandalous  but  irrefutable  conse- 
quence of  postulating  a  material  world  without  qualities  and 
a  world  of  minds  that  lack  spatiality  and  exists — nowhere^  ^^ 
One  way  of  characterizing  my  hypothesis  would  be  to  say 
that  it  is  an  effort  to  remove  this  scandal  by  showing  where- 
in the  postulation  noted  by  Dr.  Montague  is  not  true. 

The  genetic  relationships  of  my  hypothesis  can  be  still 
farther  indicated  by  coming  on  down  from  Descartes  to 
Hume  then  from  Hume  to  Huxley  and  finally  to  G.  F.  Stout 
and  John  Dewey  as  philosophers  of  to-day.     Hume's  nom- 


Sketch  of  ail  Organismal  Theory  of  Cojuciou.sn^'ss     290 

enclature  for  the  subjective  and  objective  sides  of  man's 
psychic  Hfe  is  "Relations  of  Ideas"  for  the  first,  and  '*Mat- 
ters  of  Fact"  for  the  second.  Of  the  first  kind  says  Hume, 
"Are  the  sciences  of  Geometry,  Algebra  and  Arithmetic;  and 
in  short,  every  afl^rmation  which  is  intuitively  or  demonstra- 
tively certain."  .  .  .  ''That  three  times  five  is  equal  to  half 
of  thirty/'  is  a  simple  illustration  of  the  relation  of 
ideas.  And,  "Propositions  of  this  kind  are  dis- 
coverable by  the  mere  operation  of  thought,  with- 
out dependence  on  what  is  anywhere  existent  in  the  uni- 
verse." ^^  And  further  on,  Part  2,  same  section,  we  read: 
"It  must  certainly  be  allowed,  that  nature  has  kept  us  at  a 
great  distance  from  all  her  secrets,  and  has  afforded  us  only 
the  knowledge  of  a  few  superficial  qualities  of  objects;  while 
she  conceals  from  us  those  powers  and  principles  on  which 
the  influence  of  those  objects  entirely  depends."  Then  Hume 
goes  into  a  discussion  of  the  operations  and  relations  of  the 
"superficial  qualities"  and  "secret"  powers  of  objects  which 
is  so  similar  to  my  treatment  of  the  relation  of  the  organism 
to  the  attributes  of  certain  objects  (chapters  20  and  21  this 
book,  and,  more  particularly,  my  essay  Is  Nature  luijuite?'~^ ) 
that  it  seems  as  though  his  words  must  have  been  in  my  mind 
when  I  thought  out  what  I  have  there  written,  though  I  cer- 
tainly was  not  conscious  of  Hume's  views.  And  this  sub- 
conscious influence  appears  the  more  probable  in  that  I  have 
almost  conclusive  proof  of  having  read  his  argument  not  long 
before  my  own  was  written.  I  am  certain,  however,  that  if 
his  statements  were  in  my  mind  they  Avere  only  in  its  pro- 
conscious  part  and  were  not  nor  ever  had  been  in  its  full- 
conscious  part.  In  other  words,  if  I  had  rcafl  his  words  I 
had  not  grasped  their  full  significance.  This  probable  in- 
stance of  the  "sub-"  or  "pro"-conscious  I  refer  to  not  so 
much  because  of  its  interest  in  this  instance,  as  because  of  its 
bearing  on  my  conception  of  the  nature  of  consciousness. 
The  discussion  by  Hume  to  which  I  refer  is  that  in  which  he 


300  The  Unity  of  the  Organism 

talks  about  the  sensible  qualities  and  the  "secret  powers" 
of  the  bread  we  eat.     "Our  senses  inform  us  of  the  color, 
weight,  and  consistence  of  the  bread,"  he  says,  "but  neither 
sense  nor  reason  can  ever  inform  us  of  those  qualities  which 
fit  it  for  the  nourishment  and  support  of  a  human  body." 
The    particular    puzzle    upon    which    Hume    comes    in    this 
matter  is  the  fact  that  although  the  examination  here  and 
now  of  a  natural  object  gives  us  absolutely  no  clue  as  to  what 
latent  attributes   ("secret  powers,"  he  calls  them)   the  ob- 
ject may  possess,  when  we  examine  a  second  object  of  the 
same  kind  we  assume  that  the  same  secret  powers  are  pos- 
sessed by  the  second  object.     "If  a  body  of  like  colour  and 
consistence  with  that   bread,  which  we   have   formerly   eat, 
be  presented  to  us,  we  make  no  scruple  of  repeating  the  ex- 
periment, and  foresee,  with  certainty,  like  nourishment  and 
support.     Now  this  is  a  process  of  the  mind,  of  thought," 
Hume  goes  on  to  say,  "of  which  I  would  willingly  know  the 
foundation."      "The   bread,"    he   says,    a   little    farther    on, 
"which  fonnerly  I  eat,  nourished  me ;  that  is,  a  body  of  such 
sensible  qualities  was,  at  that  time,  endued  with  such  secret 
powers :  but  does  it  follow  that  other  bread  must  also  nour- 
ish me  at  another  time,  and  that  like  sensible  qualities  must 
always   be   attended   with   like   secret   powers?     The   conse- 
quences seem  nowise  necessary.    At  least,  it  must  be  acknowl- 
edged that  there  is  here  a  consequence  drawn  by  the  mind; 
that  there  is  a  certain  step  taken;  a  process  of  thought, 
and  an  inference,  which  wants  to  be  explained."     Then  after 
a  little  further  argument  to  show  the  necessity  of  recog- 
nizing such  a  process  we  find  this  to  me  exceedingly  interest- 
ing passage:     "There  is  required  a  medium,  which  may  en- 
able the  mind  to  draw  such  an  inference,  if  indeed  it  be  drawn 
by  reasoning  and  argument.     What  that  medium  is,  I  must 
confess,  passes  my  comprehension;  and  it  is  incumbent  on 
those  to  produce  it,  who  assert  that  it  really  exists,  and  is 
the  origin  of  all  our  conclusions  concerning  matter  of  fact." 


Sketch  of  an  Orgamsmal  Theorij  of  Consciousness     1501 

The  great  merit  liere  shown  by  Hume  is  liis  ability  to  j)iish 
tlie  analysis  of  his  problem  to  the  very  hmit  of  the  positive 
information  lie  had  to  go  on,  recognise  exactly  wherein  his 
information  was  lacking,  and  then  stop  without  running  off 
into  a  purely  speculative  substitute  for  his  deficient  knowl- 
edge. According  to  my  hypothesis  the  unknown  "medium" 
which  he  saw  must  exist,  the  researches  of  a  century  and  a 
half  since  he  wrote,  in  chemistry,  physiology,  general  zoology 
and  botany,  and  psychology,  have  enabled  us  to  see  is  the 
individual  animal  organism  reaching  with  the  respiratory 
substance  (oxygen?)  it  takes  in.  In  this  one  particular  and, 
from  the  standpoint  to  which  we  have  been  accustomed,  very 
peculiar  case,  the  reaction  is  at  one  and  the  same  time  part 
of  the  essence  of  both  ideas  and  impressions  in  the  Humean 
sense,  the  reaction  being  the  "medium"  or  the  "certain  step" 
by  which  the  inference  is  drawn,  this  inferring  being  possible 
because  of  the  continuity  of  the  organism  as  a  person,  or 
self,  and  the  persistence  of  the  respiratory  substance  as  the 
same  identical  thing  from  the  past  through  the  present  into 
the  future. 

We  will  now  notice  how  Huxley,  because  of  his  much  more 
extensive  knowledge  of  the  structure  and  function  of  animals 
than  Hume  possessed,  was  able  to  draw  still  closer  than  Hume 
could  to  the  heart  of  the  old  Mind-Body  puzzle.  The  gist 
of  Huxley's  position  on,  and  contribution  to,  the  problem 
can  conveniently  be  presented  through  his  remarks  on  the 
question  of  innateness  of  various  aspects  of  psychic  life, 
these  remarks  occurring  in  his  essay  on  Hume.  After  point- 
ing out  that  neither  Locke  nor  Hume  seemed  to  know  exact- 
ly what  Descartes,  the  originator  of  the  modern  conception 
of  innate  ideas,  meant  by  his  phrase  "idees  naturelles,"  Hux- 
ley quotes  Descartes  as  follows:  "I  have  used  this  term  in 
the  same  sense  as  when  we  say  that  generosity  is  innate  in 
certain  families;  or  that  certain  maladies  such  as  gout  or 
gravel,  arc  innate  in  others ;  not  that  children  born  in  these 


302  The  Unity  of  the  Organism 

families  are  troubled  with  such  diseases  in  their  mother's 
womb;  but  because  they  are  born  with  the  disposition  or 
faculty  of  contracting  them."^^  ^}^q^  ^fter  further  quota- 
tions to  the  same  effect  Huxley  writes:  "Whoever  denies 
what  isj  in  fact,  an  inconceivable  proposition,  that  sensations 
pass,  as  such,  from  the  external  world  into  the  mind,  must 
admit  the  conclusion  here  laid  down  by  Descartes,  that, 
strictly  speaking,  sensations,  and  a  fortiori,  all  the  other 
contents  of  the  mind,  are  innate.  Or,  to  state  the  matter  in 
accordance  with  views  previously  expounded,  that  they  are 
products  of  the  inherent  properties  of  the  thinking  organ,  in 
which  they  lie  potentially,  before  they  are  called  into  exist- 
ence by  their  appropriate  causes." 

The  upshot  of  this  clearly  is  that  innate  for  Descartes  and 
Huxley  means  hardly  anything  else  than  hereditary,  as  ap- 
plied to  the  psychical  as  well  as  to  the  physical  attributes  of 
animals.  The  ample  justification  in  our  day  of  the  view 
that  psychical  attributes  are  hereditary  should,  it  would 
seem,  restore  to  full  standing  in  biology,  the  conception  of 
innate  ideas — only,  of  course,  in  a  very  different  sense  from 
that  into  which  later  Idealists  have  perverted  it. 

It  is  in  this  discussion  that  Huxley  makes  one  of  the  most 
direct  and  unanswerable  arguments  against  materialism  that 
can  be  made:  "The  more  completely  the  materialistic  posi- 
tion is  admitted,  the  easier  it  is  to  show  that  the  idealistic 
position  is  unassailable,  if  the  idealist  confines  himself  with- 
in the  limits  of  positive  knowledge."^^  That  is  to  say,  if  the 
materialist  insists  that  all  traces  of  innateness  of  ideas  and 
other  contents  of  the  mind  must  be  repudiated,  he  virtually 
contends  that  heredity  of  whatever  sort,  whether  of  physical 
or  psychical  attributes,  must  be  repudiated.  With  this  con- 
ception of  innateness  in  the  entire  psychic  aspect  of  the 
organism  before  him  Huxley  asks:  "What  is  meant  by  ex- 
perience .^" 

"It  is  the  conversion,"  he  replies,  "by  unknown  causes,  of 


Sketch   of  an  Organismal  Thcorij  of  Conscwmncss     3015 

these  innate  potentialities  Into  actual  experiences."-^  Now 
these  "unknown  causes"  are,  according  to  my  view,  essential- 
ly the  same  as  the  "medium"  which  Hume  recognized  must 
exist  for  making  the  "stej)"  i)ossll)le  from  tlie  "sujjerficial 
qualities"  to  the  "secret  j)owers"  of  natural  objects  and  from 
the  "secret  powers"  of  one  object  to  those  of  another.  They 
are,  to  repeat,  the  reaction  of  the  orrranism  in  its  latently 
psychical  aspect,  with  "the  breath  of  life,"  that  is,  with  the 
oxygen,  or  whatever  be  the  gaseous  constituent  of  the  air 
which  is  active  in  respiration.  And  I  believe  we  can  see  to 
a  considerable  extent  why  Huxley  considered  these  causes  as 
wholly  unknown.  It  was  because  physiology  and  bio-chemis- 
try in  his  day  w^ere  not  yet  able  to  view  the  organism  from 
the  standpoint  of  physical  chemistry.  Because  of  this  ina- 
bility Huxley  nor  any  other  physiologist  of  his  period  had 
an  adequate  structural  ground-work  for  thinking  organis- 
mall}^  about  living  things.  They  were  consequently  obliged, 
really,  to  think  of  all  psychic  phenomena,  and  consciousness 
with  the  rest,  as  being  restricted  to  the  nervous  system. 
That  such  was  Huxley's  view  at  any  rate,  we  know  from 
his  own  words :  "No  one  who  is  cognisant  of  the  facts  of  the 
case  nowadays  doubts,"  he  writes,  "that  the  roots  of  psychol- 
ogy lie  in  the  physiology  of  the  nervous  system."  The  im- 
portant revision  of  this  statement  which  our  hypothesis  calls 
for  is  that  while  the  roots  of  psychology  are  indeed  in  the 
nervous  system  they  are  by  no  means  in  that  system  alone. 
They  pass  through  it  to  a  much  deeper  level,  so  to  speak,  and 
in  passing  draw  great  nutriment  from  it. 

In  a  brief  but  important  ])a])er  starting  off  with  the  proj)- 
osition  that  a  philosopher  can  not  legitimately  question  the 
existence  of  the  external  world — that  all  he  can  rightly  do  is 
to  inquire  what  that  world  Is  and  how  we  can  know  it  at  all, 
G.  F.  Stout  comes  to  the  kernel  of  the  |)r()])len)  in  considera- 
bly the  same  way  that  Hume  and  Huxley  came  to  it.  "I'or 
primitive   consciousness    and    for    our   own    unreflective    con- 


304  The   Unity  of  the  Organism 

sciousness,"  he  says,  "sense  experience  and  the  correlative 
agency  which  conditions  it  coalesce  in  one  unanalysed  total 
object.  They  coalesce  in  such  a  way  that  the  sense-presenta- 
tion appears  as  possessing  the  independence  of  the  not-self, 
and  the  independent  not-self  seems  to  be  given  with  the  same 
immediacy  as  the  sense-presentation."  And,  "this  complex 
but  unanalysed  cognition,"  Stout  continues,  "is  the  germ 
from  which  our  detailed  knowledge  of  matter  develops. "^^ 
If  proved  true  my  hypothesis  would  be  a  considerable  for- 
ward step,  I  believe,  in  analysing  this  "unanalysed  cogni- 
tion." For  although  Stout's  assertion  "the  independent  not- 
self  is  not  matter"  seems  at  first  sight  to  exclude  oxygen  or 
any  other  constituent  of  our  breath  from  such  a  place  in 
the  external  world  of  his  conception  as  that  which  it  has  in 
that  world  according  to  my  conception  this  exclusion  is,  I 
think,  only  seemingly  so,  for  a  sentence  farther  on  the  author 
says  matter  "essentially  includes  the  qualification  of  the  in- 
dependent not-self  by  the  content  of  sense-experience."  The 
seeming  discrepancy  is  probably  due  to  the  generality  of  the  i 
term  matter.  I  too  would  say  that  the  "independent  not- 
self"  is  not  matter  were  I  to  mean  b}^  matter  the  total  sub- 
stance of  the  external  world.  But  in  the  sense  that  the  effec- 
tive respiratory  gas  (oxygen  supposedly)  is  matter,  my 
hypothesis  would  require  me  to  hold  that  the  not-self  has  an 
essential  material  component,  which  component  is  really  the 
attribute  of  the  gas  in  virtue  of  which  it  reacts  with  the 
organism  in  the  peculiar  way  it  does  to  produce  conscious- 
ness. It  seems  to  me  that  what  Stout  seeks  in  the  "quali- 
fication of  the  independent  not-self  by  the  content  of  sense- 
experience"  is  the  immediately  consciousness-producing  attri- 
bute of  the  respiratory  gas.  We  might  state  the  point  this 
way:  Oxygen  (or  the  effective  respiratory  gas)  has  a  double 
status  in  human  consciousness.  First  and  most  fundamental- 
ly, it  has  the  status  of  an  immediate  and  essential  participant 
in  producing  all  consciousness  whatever;  and  second  it  has 


Sketch  of  an  Orgatiismal  Thvarjf  of  Consciousness     305 

the  status  of  an  indirect  participant  in  })ro(]ucin^r  tlu>  par- 
ticular consciousness  which  we  call  observational  knowledge 
of  the  gas.  Our  knowledge  of  this  one  gas  is  due  to  two 
things,  (1)  to  our  reaction  to  it  through  our  sense  organs 
in  the  usual  psychological  meaning  of  react;  and  (ii)  to  our 
reaction  with  it  through  the  [)rotoplasniic  basis  of  all  con- 
sciousness, reaction  in  this  case  having  the  meaning  which 
chemistry  has  given  the  word.  What  the  relation  is  \m- 
tween  the  attributes  of  the  gas  in  virtue  of  which  it  reacts 
with  the  organism  in  these  two  ways,  and  also  what  the  rela- 
tion is  between  the  attributes  of  the  organism  in  virtue  of 
which  it  reacts  with  the  gas  in  these  two  ways,  are  questions 
with  which  a  theory  of  knowledxre  would  deal  but  which  lies 
outside  of  the  scope  of  this  sketch,  which,  as  has  already 
been  said,  restricts  itself  to,  a  theory  of  consciousness.  I 
may,  however,  refer  in  passing  to  the  fact  that  chemistry 
appears  to  be  all  at  sea  on  the  problem  of  the  relation  be- 
tween the  chemical  and  the  physical  attributes  of  all  sub- 
stances whatever;  so  the  difficulties  about  oxygen  in  this  one 
particular  are  not  an  unshared  difficulty. 

Finall}',  to  bring  this  exposition  of  the  historical  setting 
of  my  h^^pothesis  down  to  the  present  hour,  I  call  attention 
to  the  way  the  hypothesis  connects  with  the  best  that  formal 
philosoph}'^  in  our  own  day  has  done,  or  as  I  suspect  is 
competent  to  do,  towards  making  out  what  "experience"  is. 
No  philosopher  with  whom  I  have  met  has  gone  farther  in 
this  direction  than  John  Dewev.  In  his  recent  essav,  ./  Uc- 
covery  of  Philosophy,  we  read:  ''Dialectic  developments  of 
the  notion  of  self-preservation,  of  the  conuius  cssctidi,  often 
ignore  all  the  important  facts  of  the  actual  process.  "I'lu-y 
argue  as  if  self-control,  self-development,  went  on  directly  as 
a  sort  of  unrolling  push  from  within.  But  life  endures  only 
in  virtue  of  the  support  of  the  enriromnent.'''''^  The  italics 
are  mine  and  mark  the  most  vital  })art  of  the  quotation  for 
us.      And   a   page   farther   on:      ''Kx])erience    is    wo   sll])ping 


306  The  Unity  of  the  Organism 

along  in  a  path  fixed  by  inner  consciousness.  Private  con- 
sciousness is  an  incidental  outcome  of  experience  of  a  vitally 
objective  sort;  it  is  not  its  source.  Undergoing,  however, 
is  never  mere  passivity.  The  most  patient  patient  is  more 
than  a  receptor.  He  is  also  an  agent — a  reactor."  .  .  . 
Again  the  itahcs  are  mine.  I  take  the  liberty  to  end  the  quo- 
tation at  "reactor"  though  the  remaining  part  of  the  sen- 
tence is  important  for  Dewey's  particular  purpose.  But  my 
aim  is  different.  I  want  to  fix  attention  on  the  two  state- 
ments italicised  for  the  purpose  of  showing  how  my  hypo- 
thesis connects  with  Dewey's  general  conception  of  experi- 
ence. When  Dewey  says  life  endures  only  as  supported  by 
the  environment,  he  is  speaking  in  very  general  terms,  having 
reference,  I  imagine,  more  to  social  and  other  bulk  aspects 
of  environment.  My  hypothesis,  on  the  contrary,  makes  the 
dependence  of  life  on  environment  exceedingly  specific  in  that 
it  undertakes  to  show  the  particular  thing  in  the  environ- 
ment, namely,  the  respiratory  part  of  the  atmosphere,  which 
is  physiologically  basal  to  self-development  and  self-pre- 
servation. The  Self  which  traditional  philosophy  has  strug- 
gled so  hard  to  understand  is  literally,  the  human  organism, 
according  to  my  hypothesis.  And  when  in  this  discussion  I 
speak  of  it  as  reacting  with  the  respiratory  air  to  produce 
consciousness,  I  am  using  the  verb  to  react  in  a  very  specific, 
physico-chemico-biological  sense,  while  Dewey  is  using  it  in 
a  general  sense,  and  explicitly  at  least,  with  only  a  psy- 
chological implication. 

The  "self"  which  I  am  suggesting  does  indeed  imply 
"another"  no  less  unequivocally  than  does  the  "self"  of  ad- 
vanced social  psychology.  But  the  "self"  and  the  "other" 
implied  by  my  hj^pothesis  differ  from  those  of  current  philo- 
sophical theory  in  that  the  roots  of  both  are  not  only  in 
the  social  relationships  of  the  human  species,  but  extend 
right  on  through  these  into  sub-human  relationships,  even 
down  into  the  very  constitution  of  inorganic  nature.     The 


Sketch  of  an  Organismal  Thcorij  of  Con.srionstwss     807 

"self"  and  the  "other"  of  my  conception  are  more  per.sonallv 
objective,  and  more  cosmic  in  their  affinities,  than  arc  tiie 
"self"  and  the  "other"  of  social  psychology. 

Continuing  now  with  our  examination  of  the  foundation 
of  my  hypothesis  I  find  it  convenient,  especially  because  of 
my  reference  a  few  pages  back,  to  Huxley's  unanswerable 
contention  for  an  essence  of  truth  in  both  materialism  and 
idealism,  to  call  attention  to  a  natur.il  history  fact  in  the 
higher  mental  life  of  man  which  I  take  to  be  a  strong  con- 
firmation of  the  contention.  This  fact  concerns  the  general 
difference  between  what  are  commonly  known  as  the  mate- 
rialistic and  the  idealistic  attitudes  of  mind.  This  difference 
comes,  I  believe,  to  the  same  thing  finally,  as  the  difference 
between  the  objective  and  subjective  attitudes,  and  is  also 
the  difference,  at  bottom,  between  what  in  ratlier  loose  thouifh 
prevalent  expression,  is  called  the  difference  between  the 
scientific  and  the  philosophic  attitudes.  It  would  seem  that 
the  philosopher  who  declares  himself  to  be  an  Absolute  Ideal- 
ist, as  Royce  does,  is  under  heavy  obligation,  especially  if 
he  enters  the  field  of  psychology,  to  explain  the  fact  that  the 
originators  of  great  interpretative  ideas  of  nature  have  in- 
variably recognized  that  their  hypotheses  must  be  "proved'"; 
that  is,  that  the  subjective  experience  which  constitutes  the 
hypothesis  must  be  found  to  have  its  counterpart  in  tlie  ex- 
ternal world  of  sense.  If  "Reason  creates  the  world,"  even 
in  the  recondite  meaning  of  Royce's  philoso})hy,  how  hap- 
pened it  that  Newton  should  have  been  so  "restless"  for  evi- 
dence of  an  objective,  an  external  counterpart  to  the  subjec- 
tive result  he  had  reached  by  mathematical  reasoning,  that 
he  held  back  liis  reasoned  creation  for  sixteen  years,  waiting 
for  the  proof,  the  sense-perceptual  or  at  least  the  sense- 
-perccj)tible  experience,  that  should  round  out  his  reasoned 
truth .'^  May  not,  I  ask,  the  very  kernel  of  the  difference 
between  science  at  its  best  and  j)lii]os()|)hy  at  its  l)est  Ik-  in 
this,  that  the  typical  scientist  is  somewhat  deficient  in  "nst- 


308  The  Unity  of  the  Organism 

lessness,"  adopting  Royce's  terminology,  for  internal  or  sub- 
jective reality;  while  the  philosopher  of  the  schools  is  some- 
what deficient  in  restlessness  for  external  or  objective  reality? 
We  could  say  with  almost  literal  chemical  accuracy  that  the 
curiosity  and  eagerness  of  the  naturalist  for  yet  unobserved 
objective  truth  is  due  to  an  unsatisfied  affinity  which  is  weak, 
or  in  some  instances,  wholly  lacking,  in  the  subjective  idealist. 
The  facts  which  seem  to  justify  our  chemico-organismal 
hypothesis  of  conscious  psychic  life,  seem  abo  to  imply  a 
complete  interpenetration  of  objective  science  and  idealistic 
philosophy. 

As  to  the  Lowest  Terms  of  Self -Consciousness 

Let  us  now  veer  our  course  in  examining  self-conscious  life, 
and  see  what  can  be  made  out  about  its  roots  and  rootlets 
instead  of  about  its  fruitage. 

We  are  often  reminded  that  our  knowledge  about  our  in- 
ternal organs,  our  heart,  liver,  lungs,  et  cetera,  comes  only 
through  observations  by  the  anatomist  and  physiologist; 
that  we  are  quite  unconscious  of  these  organs  in  our  own 
bodies,  especially  if  they  are  working  normally.  Now  I 
point  out  that  to  be  perceptually  conscious  of  a  liver,  let  us 
say,  as  a  specialized  morphological  entity  performing  its 
appropriate  functions,  is  a  very  different  matter  from  being 
conscious  of  those  primal,  undifferentiated  processes  which 
are  basal  to  life  itself,  and  so  are  common  to  all  the  tissues 
whether  liver,  muscle,  brain,  or  what  not,  so  long  as  they  are 
actually  living.  That  that  which  is  truly  organic,  in  the 
sense  of  pertaining  to  the  fully  constituted  organism,  must 
be  regarded  from  this  standpoint  as  well  as  from  the  stand- 
point of  their  final  state  of  differentiation,  is  one  of  the 
common-places  of  modern  biology.  Let  a  person  in  as  near- 
ly perfect  health  as  he  ever  experiences,  do  his  best  to  elimi- 
nate   all    external    and    internal    stimuli    of   his    specialized 


Sketch  of  an  Organi.sfna]  Tlicorij  of  Consciousness     309 

sensory  parts;  also  all  renieinbcring,  all  ficlin<r  of  the  usual 
kind,  all  imagining,  and  all  thinking.  TIrii  Kt  him  answer 
the  question:  How  do  1  know  I  am  alive?  An  undertaking 
of  this  sort  is  wholly  introspective  in  the  sense  of  Ixing 
such  that  each  j)erson  nmst  engage  in  it  for  himself  alone. 
Pie  can  not  show  his  results  to  anybody  else.  A  good  bit  ot 
ingenuity  may  be  exercised  on  it  and  the  outcome  will  \)v 
found  to  be  rather  surprising  if  not  very  conclusive  as  to  the 
purpose  for  which  the  experiment  was  tried.  But  the  results 
as  reported  may  be  of  some  value.  Personally,  1  believe  I 
can  follow  my  consciousness  down  to  where  I  can  recognize 
its  most  basal  remaining  "content"  to  be  an  awareness  of 
what  I  may  call  extension  without  definite  limitations.  It 
seems  to  me  I  can  detect  something  to  whicli  I  could  not, 
from  its  nature  alone,  apply  the  terms  "I"  or  "me"  as  some- 
thing differentiated  from  everything  else.  Possibly  what  I 
note  is  wholly  fanciful,  but  I  seem  to  feel  myself  in  about 
the  condition  of  psychical  life  which  I  imagine  a  star  fish  is  in. 
Of  course  I  realize  how  far  such  a  statement  is  from  being 
purified  of  all  thought  and  other  ordinary  mental  elements. 
Nevertheless,  I  believe  it  to  be  of  some  value  as  evidence 
that  consciousness  is  an  attribute  of  the  organism  as  a 
whole,  and  can  neither  be  held  to  contain  an  element  which 
can  exist  separately  from  the  organism,  nor  be  restricted 
to  any  particular  part  of  the  organism  as  the  brain  or  the 
nervous  system.  There  seems  to  be  som£  evidence  "directly 
felt  by  us  ourselves,"  and  that  evidence  points  to  this  con- 
clusion as  to  the  nature  and  "seat"  of  consciousness.  The 
point  is  susceptible,  I  am  quite  sure,  of  rather  rigid  experi- 
mental examination.  However,  the  further  experiments 
which  have  suggested  themselves  to  me  involve  difficulties 
more   formidable  than   I  have   thus   far  been   in   ])osltion   to 

attempt. 

The  reader  acquainted  with  James's  notable  Chai)ter  X, 
"The  Consciousness  of  Self"  (The  Principles  of  rsjichohum. 


310  The  Unity  of  the  Organism 

Vol.  1)  will  recognize  the  difference  between  such  introspec- 
tive experimentation  as  that  here  indicated,  and  that  so  il- 
luminatingly  described  by  James  as  tried  on  himself.  While 
James's  undertaking  was  to  give  an  account  of  the  thought 
and  other  processes  in  consciousness  as  he  could  observe  them 
in  himself,  what  I  want  to  accomplish  requires  me  to  get  rid 
of,  to  ignore  as  far  as  possible,  the  very  things  which  James 
was  studying.  I  want  to  find  whether  any  "content  of  con- 
sciousness" remains  after  thought  and  the  other  usual  men- 
tal contents  are  out  of  the  reckoning.  I  believe,  however, 
that  James  opens  the  way  to  such  an  hypothesis  as  mine. 
Thus  in  a  footnote  we  read,  "The  sense  of  my  bodily  exist- 
ence, however  obscurely  recognized  as  such,  may  then  be  the 
absolute  original  of  my  conscious  selfhood,  tlie  fundamental 
perception  that  /  am.  All  appropriations  may  be  made  to 
it  hy  a  Thought  not  at  the  moment  immediately  cognized  by 
itself.  Whether  these  are  not  only  logical  possibilities  but 
actual  facts  is  something  not  yet  dogmatically  decided  in 
the  text."^ 

Except  for  a  little  misgiving  arising  from  uncertainty  as 
to  the  exact  meaning  of  "Thought"  in  this  quotation,  I  be- 
lieve my  hypothesis  does  what  James  says  his  text  leaves  un- 
decided. 

This  foot-note  of  James's  may  serve  as  a  switch  key  to 
shift  the  current  of  our  discussion  from  the  psycho-con- 
scious phase  of  life  through  the  psycho-physical  to  the  purely 
physico-chemical  phase.  The  course  along  which  this  shifting 
will  run  can  be  designated  thus :  full-fledged  intellect  ( al- 
ready examined),  instinct,  emotion,  bio-physico-chemical  or- 
ganization. 

Instinct  and  Physical  Organization 

The  discussion  from  which  we  have  just  turned  of  the 
relation  between  "inner"  and  "outer,"  between  "subjective" 


SKetch   of  an  Orgduismal  Theory  of  Conaciousmss     ,S11 

find  "objective,"  must  be  regarded  as  iiR'etiii«r  the  reciuire- 
nients  of  this  sketch  so  far  as  the  first  iiuiul><.r  ui'  tlic  series 
is  concerned;  and  the  relation  between  instinct  and  physical 
organization  will  now  receive  attention.  Tlie  evidence  of 
vital  connection  here  is  so  abundant  and  clear-cut,  and  the 
views  of  competent  observers  are  so  unanimous  tliat  tlie  sul>- 
ject  can  be  disposed  of  quite  sunnnarily.  Probably  the  most 
indubitable  single  block  of  evidence  comes  from  nest-buihbng 
and  cocoon-spinning  insects.  Many  of  the  facts  from  this 
field  have  been  so  much  ex])l()ited  for  the  very  })urposes  to 
which  we  now  invoke  them  that  a  few  quotations  from  and 
remarks  upon  the  writings  of  naturalists  generally  acknowl- 
edged for  learning  and  judicious  tliinking  will  suffice. 

We  turn  first  to  W.  M.  Wheeler,  and  take  to  begin  with, 
words  w^hich  he  in  turn  quotes  from  Bergson  :  ''As  Jkrgson 
says,"  we  read,  "  "It  has  often  been  remarked  that  most  in- 
stincts are  the  prolongation,  or  better,  the  achievement,  of 
the  work  of  organization  itself.  Where  does  the  activity  of 
instinct  begin .^^  Where  does  that  of  nature  end?  It  is  im- 
possible to  say.  In  the  metamorphoses  of  the  larva  into  the 
nymph  and  into  the  perfect  insect,  metamorphoses  which 
often  require  appropriate  adaj^tations  and  a  kind  of  initia- 
tive on  the  part  of  the  larva,  there  is  no  sharp  line  of  de- 
marcation between  the  instinct  of  the  animal  and  the  organiz- 
ing work  of  the  living  matter.  It  is  immaterial  whetlier  wc 
say  that  instinct  organizes  the  instruments  which  it  is  going 
to  use,  or  that  the  organization  })rolongs  itself  into  the  in- 
stinct by  which  it  is  to  be  used.'  "  And  Wheeler  continues: 
"The  spinning  of  the  cocoon  by  the  larval  ant  is  a  good 
example  of  the  kind  of  instinct  to  which  Bergson  refers. 
From  one  point  of  view  this  is  merely  a!i  act  of  development, 
and  the  cocoon,  or  result  of  the  secretive  activity  of  the  seric- 
teries  and  of  the  spinning  movements  of  the  larva,  is  a  pro- 
tective envelope.  But  an  enveloi)e  with  the  same  protective 
function  may  be  produced  by  other  iioeet   larv.e  >iiiiply  as  a 


312  The  Unity  of  the  Organism 

thick,  chitinous  secretion  from  the  whole  outer  surface  of  the 
hjpodermis.  Here,  too,  we  have  an  activity  which,  though 
manifested  in  a  very  different  way,  is  even  more  clearly  one 
of  growth  and  development.  And  when  the  workers  of 
(Ecophylla  or  Polyrhachis  use  their  larvae  for  weaving  the 
silken  envelope  of  the  nest,  as  described  in  Chapter  XIII, 
we  have  a  further  extension  and  modification  of  the  cocoon- 
spinning  activities.  In  this  case  the  spinning  powers  of  the 
larva  are  utilized  for  the  purpose  of  producing  an  envelope, 
not  for  its  individual  self,  but  for  the  whole  colony.  In 
conventional  works  this  latter  activity  would  be  assigned  a 
prominent  place  as  a  typical  instinct,  the  spinning  of  the 
cocoon  might  also  be  included  under  this  head,  but  the  form- 
ation of  the  puparium,  or  pupal  skin,  would  be  excluded 
as  a  purely  physiological  or  developmental  process,  yet  this 
last,  no  less  than  the  two  other  cases,  has  all  the  fundamental 
characteristics  of  an  instinct."^ 

Then  immediately  follows  this  statement,  especially  signi- 
ficant for  the  proposition  of  our  hypothesis  which  assigns  to 
the  individual  organism  the  chemical  value  of  an  elementary 
substance:  "Viewed  in  this  light  there  is  nothing  surprising 
about  the  complexity  and  relative  fixity  of  an  instinct,  for  it 
is  inseparably  correlated  with  the  structural  organization, 
and  in  this  we  have  long  been  familiar,  both  with  the  de- 
pendence of  the  complexity  and  fixity  of  parts  on  heredity 
and  the  modifiability  of  these  parts  during  the  life-cycle 
of  the  individual.  Fixed  or  instinctive  behavior  has  its 
counterpart  in  inherited  morphological  structure  as  does 
modifiable,  or  plastic,  behavior  in  well-known  ontogenetic 
and  functional  changes." 

The  statement  that  surprise  is  largely  taken  away  from 
such  elaborate  manifestations  of  instinct  as  those  here  de- 
picted, by  recognizing  that  the  instincts  are  "inseparably 
correlated  with  structural  organization"  and  have  their 
"counterpart  in  inherited  morphological  structure,"  will,  no 


Sketch  of  an  Organismal  Theory  of  Consciousness     31  Ji 

doubt,  receive  the  assent  of  most  zoologists,  as  will  also  the 
statement  that  our  lonf^  familiarity  with  structural  organi- 
zation and  morphological  inheritance  is  what  makes  us  re- 
gard these  without  surprise,  and,  by  inference,  as  compre- 
hensible. It  is  not  that  the  corporeal  form  and  structure  of 
the  worker  ants  and  of  the  larvae  which  they  manipulate  as 
spinning  instruments  and  shuttles  for  making  the  nest,  are 
necessarily  simpler  and,  on  that  account,  more  com])rehen- 
sible  than  are  the  instinctive  acts  of  the  workers,  but  that 
during  our  whole  lives  we  have  been  familiar  with  structure, 
and  ourselves  exist  as  "structural  organizations."  Tliis  is 
equivalent  to  saying  that  we  have  always  been  not  only  learn- 
ing but  directl}^  experiencing  interdependences  and  correla- 
tions among  the  common  body-parts  and  body-acts,  and  so 
regard  them  as  comprehensible,  as  explicable.  To  compre- 
hend really  an  external  complex  of  structures  and  activities 
is  to  live  the  counterpart  of  it.  To  understand  such  a  com- 
plex scientifically  is  to  understand  it  through  a  course  of 
observation  and  reasoning;  that  is,  rationally.  To  explain 
such  a  complex  is  to  bring  in,  or  recognize  consciously  one  by 
one  the  constituent  elements  of  the  complex,  and  recognize 
all  these  as  parts  of  the  ens£mhle.  It  is  to  recognize  the 
elements  in  both  their  isolate  and  integrate  capacities. 

So  much  for  the  evidence  of  integration  between  instinct 
and  physical  organization  as  presented  by  one  carefully  i)hil- 
osophical  naturalist.  Several  other  naturalists  have  gone 
nearly  as  far,  but  this  single  instance  is  so  typical  and  conclu- 
sive as  to  the  objective  facts  that  it  will  suffice.  In  com- 
menting on  the  significance  of  being  surprised  at  such  rarely 
witnessed  performances  as  those  furnished  by  these  ants, 
w^hile  we  are  not  surprised  at  common  structures  and  acts 
of  equal  or  greater  complexity  furnished  by  more  familiar 
animals  and  by  ourselves,  I  go  beyond,  though  only  a  little 

beyond  Wheeler.  . 

The  only  other  zoologist  to  whom  I  turn  for  evidence  of 


314  The  Unity  of  the  Organism 

vital  relation  between  instinct  and  structure  is  C.  O.  Whit- 
man.     His   testimony    supplements   Wheeler's   in   that   it   is 
more  exclusively  and  radically  objective  than  is  Wheeler's; 
that  is,  it  verges  less  toward  the  subjective- type  of  presenta- 
tion and  draws  nearer  to  the  bio-chemical  ground  work.     Al- 
though Whitman  wrote  relatively  little  on  animal  behavior, 
that  little  seems  to  me  to  contain  some  of  the  most  important 
observations   and   conclusions   which  have   been  produced   in 
this  branch  of  zoology.     What  I  utilize  is  taken  from  his 
address  Animal  Behavior.     The  animals  upon  which  Whit- 
man's chief  studies  were  made  were  leeches  of  the  genus  Clep- 
sine;  a  salamander  (Necturus)  ;  and  pigeons  of  several  spe- 
cies.    Our  purpose  will  be  best  served  by  quoting  a  few  sen- 
tences which  go  direct  to  the  heart  of  the  question  in  hand, 
that   namely   of  the   vital   connection   of  instinct   and   basal 
physical  structure.     "The  view  here  taken,"  Whitman  writes, 
"places  the  primary  roots  of  instinct  in  the  constitutional  ac- 
tivities of  protoplasm  and  regards  instinct  in  every  stage  of 
its  evolution  as  action  depending  essentially  upon  organiza- 
tion".^    Then,  apparently  to  clarify  and  emphasize  the  last 
clause  about  the  dependence  of  instinct  or  organization,  he 
adds  a  footnote  thus :     "Professor  Loeb  refers  instinct  back 
to  '(1)  polar  differences  in  the  chemical  constitution  in  the 
Qgg  substance,  and  (2)  the  presence  of  such  substances  in  the 
^gg  as  determine  heliotropic,  chemotropic,  stereotropic,  and 
similar  phenomena  of  irritability.'     According  to  this  view, 
the  power  to  respond  to  stimuli  lies  in  unorganized  chemical 
substances,  and  the  same  powers  exist  in  the  adult  as  in  the 
Qgg^  because  the  same  chemical  substances  are  present.     Or- 
ganization serves  at  all  stages  merely  as  a  mechanical  means 
of  giving  definite  directions  to  responses. 

"The  view  I  have  taken  regards  instinctive  action  as 
organic  action,  whatever  be  the  stage  of  manifestation.  The 
Qgg  differs  from  the  adult  in  having  an  organization  of  a 
very  simple  primary  order,  and  correspondingly  simple  pow- 


Sketch  of  an  Orc/anismal  'J'/wor//  of  Conscioustwss     315 

ers  of  response.  Instinct  and  oi  .ranization  arc,  to  me,  two 
aspects  of  one  and  the  same  tl)in«r,  Ucnvv  both  have  onto- 
genetic and  pliylogenetic  development.*' 

Tiiese  statements  sliow,  as  do  those  given  in  our  disc-iission 
of  the  cell-tlieory,  how  far  Whitman  went  away  from  full- 
fledged  elenientalism  and  toward  organismalism.  Hut  his 
treatment  of  instinct  and  animal  behavior  reveals  what  his 
treatment  of  the  cell-theory  does  not,  at  least  so  clearly; 
namely,  how  far  he  also  went  on  the  way  to  the  natural  his- 
tory mode  as  contrasted  with  the  mechanistic  mode  of  j)hil- 
osophizing  on  biological  phenomena.  And  this  gives  me  a 
pleasant  opportunity  to  testify  to  the  genuinely  naturalist 
current  that  ran  through  his  life  and  work.  An  unform't table 
visit  which  I  liad  with  him  among  his  pigeons  not  long  before 
he  died,  permitted  me  to  see  something  of  the  character  and 
depth  of  his  interest  in  those  animals.  His  whole  attitude 
toward  them — his  wonderfully  broad  information  about,  and 
understanding  of  their  general  ways  of  life  and  personal 
idiosyncrasies,  his  solicitude  for  them,  and  his  measured  af- 
fection for  them — was  such  as  is  never  displayed  bv  anv 
one  who  has  not  very  much  of  the  real  naturalist  about  him, 
in  his  personality  as  well  as  in  his  knowledge.  The  individual 
pigeons,  many  of  them  at  any  rate,  appeared  to  Ix^  realities 
to  him  in  a  deep  sense  and  not  merely  "mechanical  means  for 
giving  definite  directions  to  responses"  of  chemical  sul)- 
stances.  But  after  all  this  is  said,  it  must  also  be  said  that 
there  is  no  evidence  that  Whitman  ever  grasped  fully  the  con- 
ception that  the  "constitutional  activities  of  protoplasm"  in 
which  he  believed  instincts  to  be  rooted,  nuist  l)e  the  consti- 
tutional activities  oi  protoplasms  (protoplasm  in  thi'  plural 
number),  because  no  individual  })igeon  is  either  any  other  in- 
dividual nor  even  exactlv  like  anv  other;  and  also  that  the 
existence  of  protoplasms  is  de])endent  upon  the  organisms 
to  w^hich  they  belong  as  well  as  upon  the  chemical  substances 
of  which  they  are  composed.     Whitman  went  so  far  on  tiie 


316  The  Unity  of  the  Organism 

road  toward  organismalism  as  to  believe  genuinely  in  the 
organic  and  organisation,  but  not  far  enough  to  make  him 
accept  unreservedly  individual  organisms. 

We  are  able  to  state  definitely  wherein  lies  the  great  and 
rather  unique  merit  of  Whitman's  investigations  on  animal 
behavior.  (1)  By  a  judicious  combination  of  pure  observa- 
tion and  observation  aided  by  experiment  and  conception,  he 
pushed  psychic  phenomena  in  the  form  of  instinct  down  al- 
most to  the  physico-chemical  level ;  that  is,  to  the  proto- 
plasmic level.  (2)  He  at  the  same  time  remained  positively 
within  the  organic,  the  living  realm.  His  merit  is  that  of 
restraint  as  well  as  of  positive  achievement.  He  did  not  per- 
mit his  enthusiasm  for  physical  explanation  to  betray 
him  into  adopting  a  phraseology  Avhich,  while  sounding  like 
an  explanation  of  instinct,  amounts  in  reality  to  a  denial  or 
a  repudiation  of  it. 

So  much  for  the  evidence  of  vital  connection  between  in- 
stinct and  organization.  According  to  the  schedule  indi- 
cated a  few  pages  back  for  reviewing  systematically  this  con- 
nection through  the  entire  range  of  psychic  life,  we  have 
next  to  glance  at  the  connection  between  the  emotions  and 
organization. 

Emotion  and  Physical  Organization 

Approaching  this  subject  as  we  now  are  from  the  direction 
of  psychology  proper,  the  well-known  James-Lange  interpre- 
tation of  emotion  comes  immediately  to  mind.  It  will  be 
advantageous  for  our  sketch  not  to  focus  attention  too  close- 
ly on  any  theory  or  discussion  but  to  take  in  as  much  as  we 
can  of  the  entire  field,  keeping  in  the  foreground  our  own 
personal  experiences  and  observations  as  contrasted  with  the 
descriptions  and  views  of  authorities.  What  I  mean  is  that 
the  reader  shall  take  himself  in  hand  for  serious  study  as  to 
his  emotional  life,  w^atching  himself  from  hour  to  hour,  day 


Sketch   of  an  OrganisinaJ  Theory  of  Consciousness     317 

to  day,  and  year  to  year  under  all  the  varied  conditions, 
happenings,  purposes,  and  impulses  to  which  he  is  subject. 
In  doing  this  a  special  j)oint  should  be  made  of  looking  back 
scrutinizingly  at  experiences  of  particular  satisfaction,  ela- 
tion, joy,  sorrow,  irritation,  anger,  fear,  dread,  humiliation, 
and  shame,  as  soon  after  their  occurrence  as  possible  that 
they  ma}^  be  fresh  in  memory.  But  incidents  and  episodes 
of  one's  remoter  past  which  stand  out  with  special  vividness 
from  the  intensity  of  the  particular  emotions  when  they  were 
experienced,  or  because  of  results  which  flowed  from  them, 
will  be  found  illuminatin";. 

To  what  extent  and  in  what  particular  fashion  was  our 
bodily  organization  implicated  in  the  feelings  and  emotions 
W'e  experienced,  is  our  problem.  Fortunately  one  can  "live 
over  again"  as  we  sa}- ;  can  "work  himself  into"  rather  pro- 
nounced emotional  states,  through  a  combination  of  memory 
and  imagination.  That  is,  he  can  be  much  of  a  genuine  dram- 
atist when  all  alone,  as  touching  events  and  scenes  of  his 
own  past  experience.  What  happens  to  your  body  when  you 
do  that  sort  of  thing?  is  the  central  question  before  us.  The 
very  criterion  by  which  you  answer  this  question  you  will 
find  will  be  that  of  how  far  the  body-manifestations  a})])ro- 
priate  to  the  particular  emotions  arc  elicited  through  your 
efforts.  If  your  hands  do  not  clinch  somewhat,  if  many  of 
your  arm,  leg,  and  abdominal  muscles  do  not  contract  some- 
what, if  your  respiration  does  not  cjuicken  somewhat,  and 
other  manifestations,  various  corporeal  indices  of  anger,  do 
not  appear  quite  inde})endently  of  direct  Intenticm  on  your 
part,  you  will  be  sure  you  have  not  "worked  u})''  a  geminu- 
state  of  anger.  The  only  real  knowledge  of  an  emotion  is  a 
lived  knowledge  of  that  emotion.  In  order  to  be  a  true  actor 
your  body  parts  must  act,  directly,  automatically,  s]ion- 
taneously,  so  far  as  any  rational  purpose  is  concerned.  And 
what  is  true  of  anger  is  clearly  true  of  all  other  emotions. 

Our  emotional  activities  mav   Ix'  described   as   instinctive 


318  The  Unity  of  the  Organism 

and  reflex  activities,  the  feeling-impulse  of  which  comes 
through  intelligence,  but  is  not  of  intelligence — is  not  under 
the  direct  guidance  and  control  of  intelligence.  According 
to  this  interpretation  no  animal,  no  matter  how  highly  con- 
stituted as  to  instincts  and  reflexes,  could  have  emotion  un- 
less it  had  intelligence.  Emotional  activity  is  instinctive  and 
reflex  activity  of  an  intelligent  organism,  with,  however,  the 
element  of  intellect  eliminated  or  in  abeyance  for  the  time 
being  as  regards  these  particular  acts.  This  is  what  I  would 
call  the  natural  history  description  of  emotion.  And  I  be- 
lieve it  is  in  essential  accord  with  James's  conception  of  emo- 
tion, but  his  description  is  a  psycho-physiological  rather 
than  a  natural  history  description.  I  am  quite  sure  that 
what  I  have  just  said  means  virtually  the  same  as  the  follow- 
ing: "7/  we  fancy  some  strong  emotion,  and  then  try  to  ab- 
stract from  our  consciousness  of  it  all  the  feelings  of  its 
bodily  symptoms,  we  find  we  have  nothing  left  behind,  no 
'mind-stuff*'  out  of  which  the  emotion  can  be  constituted, 
and  that  a  cold  and  neutral  state  of  intellectual  perception 
is  all  that  remains."^ 

I  will  now  point  out  wherein  I  believe  the  natural  history 
description  and  interpretation  of  emotion  are  somewhat  truer 
and  better  than  those  given  by  James  and  other  physiologi- 
cal psychologists — and,  I  may  add — very  much  truer  and 
better  than  those  given  by  certain  writers  who  approach  the 
subject  from  the  physiological  side  pure  and  simple.  James's 
epigrammatic  statements  about  being  afraid  because  we 
tremble  when  we  meet  a  bear  in  the  woods ;  about  being  sorry 
because  we  cry ;  about  being  angry  because  we  strike,  do  his 
OAvn  position  some  injustice,  I  think.  This  is  an  instance  in 
which  his  gift  for  piquant  writing  succeeded  too  well.  But 
the  fact  ought  to  be  noticed  that  what  he  actually  says  is 
that  as  between  the  usual  statement,  namely,  that  we  tremble 
because  we  are  afraid,  cry  because  we  are  sorry,  strike  be- 
cause we  are  angry,  and  his  way  of  stating  the  case,  his^way  is 


I 


Sketch  of  an  Orcjatiismal  llicorif  of  Consciousness     811) 

^^more  rational."  It  is  only  relative,  not  absolute  tnitli,  he 
is  aiming  at  in  tliese  statements.  Nevertheless,  after  due  al- 
lowance is  made  for  an  expressional  miscue  to  some  extent, 
there  is  yet  substantial  defect  in  his  ])resentation.  Speaking 
in  general  terms,  the  defectiveness  is  not  so  much  in  the 
antithesis  set  up  as  in  the  rcstrictedness  implied.  Or,  bring- 
ing the  criticism  around  toward  our  ])articular  standpoint, 
the  statement  falls  short  of  being  organismal. 

Cannon  has,  I  believe,  indicated  the  direction  in  which  the 
adequate  statement  lies.  He  writes:  "We  do  not  'feel  sorry 
because  we  cry,'  as  James  contended,  but  we  cry  Ix'canse 
when  we  are  sorry  or  overjoyed  or  violently  angry  or  full  of 
tender  affection — when  any  one  of  these  diverse  emotional 
states  is  present — there  are  nervous  discharges  by  sympathe- 
tic channels  to  various  viscera,  including  the  lachrymal 
glands.  In  terror  and  rage  and  intense  elation,  for  examj)le, 
the  responses  in  the  viscera  seem  too  uniform  to  offer  a  satis- 
factory means  of  distinguishing  states  which,  in  man  at  least, 
are  very  different  in  subjective  quality.  For  this  reason  I 
am  inclined  to  urge  that  the  visceral  changes  merelv  contri- 
bute to  an  emotional  complex  more  or  less  intlefinite,  but 
still  pertinent,  feelings  of  disturbance  in  organs  of  which 
we  are  not  usually  conscious."  ^^  What  Cannon's  criticism 
amounts  to,  expressed  in  other  language  is :  while  freely 
granting  that  organs  and  functions  in  the  usual  ])hysiologi- 
cal  sense  play  an  essential  part  in  emotion,  neither  the  vis- 
ceral nor  any  other  single  set  of  organs  is  sufficient  to  account 
for  the  whole  of  any  emotion.  Msceral  changes  contribute 
to  the  "emotional  com])lex,"  but  the  real  source  of  the  feel- 
ings involved  is  embedded  elsewhere  and  more  broadly  in  the 
organization.  Cannon  suggests:  "the  natural  response  is  a 
pattern  reaction,  like  inborn  reflexes  of  low  order."  ^^  "The 
typical  facial  and  bodily  expressions,"  he  writes,  "automati- 
cally assumed  in  different  emotions,  indicate  discharge  of  pe- 
culiar groui)ings  of  neurones  in  the  several  effective  states." 


320  The  Unity  of  the  Organism 

Without  stopping  to  examine  this  language  in  detail,  our 
aim  will  be  achieved  by  pointing  out  that  the  more  closely  the 
various  emotions  are  scrutinized,  and  the  more  effort  there 
is  made  to  refer  them  to  their  causes,  the  more  varied  are  they 
found  to  be,  and  the  more  widely  are  we  led  to  search  in  the 
organization  for  causal  factors.  The  mental  attitude  of  per- 
fect openness  toward  any  and  all  facts,  both  of  effect  and 
cause,  which  may  occur  in  a  given  organic  situation,  is  one 
of  the  leading  characterizations  of  the  organismal  conception. 
The  assertion  that  the  organism  as  a  whole  is  the  causal  ex- 
planation of  an  emotion  or  an  "emotion  complex"  is  justified 
by  two  considerations:  (1)  Except  for  the  organism  viewed 
alive  and  whole  and  under  both  its  ontogenic  and  phylogenic 
aspects,  the  emotion  would  not  exist;  and  (2)  so  wide-spread 
and  subtle  does  common  observation  recognize  the  parts  of 
the  organism  involved  to  be  in  many  of  its  emotional  activi- 
ties that  for  practical  purposes,  it  is  better  to  work  on  the 
hypothesis  that  all  parts  of  the  organism  are  implicated  than 
to  adopt  the  alternative  hypothesis  that  certain  parts  only 
are  involved;  that  is,  that  some  parts  are  not  involved. 

As  a  matter  of  fact,  I  believe  that  in  spirit  James'  hypo- 
thesis is  organismal  even  though,  probably  from  his  training 
and  career  in  formal  anatomy,  physiology,  and  psychology, 
he  never  became  entirely  free  from  the  Body-Soul  antithesis 
and  the  dogmatisms  of  "nerve  physiology,"  which  have  so 
dominated  modern  physiology  and  psychology.  This  opinion 
I  base  on  the  general  tenor  of  his  discussions  particularly 
of  the  emotions,  rather  than  on  his  direct  formulation  of  his 
theory  of  emotion.  I  will  quote  a  few  passages  that  seem 
particularly  to  trend  in  this  direction.  "No  reader  of  the 
last  two  chapters  \The  Production  of  Movement,  and  In- 
stinct^ will  be  inclined  to  doubt  the  fact  that  objects  do 
excite  bodily  changes  by  a  preorganized  mechanism,  or  the 
farther  fact  that  the  changes  are  so  indefinitely  numerous 
and  subtle  that  the  entire  organism  may  be  called  a  sound- 


Sketch  of  an  Organismal  Theory  of  Consciousness     3iil 

ing-hoard,  which  every  change  of  consciousness,  however 
slight,  may  make  reverberate.  The  various  premutations  and 
combinations  of  which  these  organic  activities  are  susceptible 
make  it  abstractly  possible  that  no  shade  of  emotion,  how- 
ever slight,  should  be  without  a  bodily  reverberation  as 
unique,  when  taken  in  its  totality,  as  is  the  mental  mood  it- 
self. The  immense  number  of  parts  modified  in  each  emotion 
is  what  makes  it  so  difficult  for  us  to  reproduce  in  cold  blood 
the  total  and  integral  ex])ression  of  any  one  of  them.  Wi- 
may  catch  the  trick  with  the  voluntary  muscles,  but  fail  witli 
the  skin,  glands,  heart,  and  other  viscera." ^^  I  ask  the  read- 
er to  make  special  note  of  the  part  of  the  quotation  be- 
ginning, ''The  various  permutations"  as  we  shall  have  more 
to  say  about  it  a  few  pages  farther  on. 

Again  we  read:  "Our  whole  cubic  capacity  is  sensibly 
alive ;  and  each  morsel  of  it  contributes  its  pulsations  of  feel- 
ing, dim  or  sharp,  pleasant,  painful,  or  dubious,  to  that  sense 
of  personality  that  every  one  of  us  unfamiliarly  carries  with 
him.  It  is  surprising  what  little  items  give  accent  to  these 
complexes  of  sensibility."^^  I  hope  the  reader  will  notice 
how  easy  it  would  be  for  me  to  contend  that  these  state- 
ments come  near  to  my  statement  about  "inner"  and  "outer," 
or  subjective  and  objective;  and  also  to  my  formal  hypo- 
thesis as  to  the  nature  of  consciousness.  However,  I  do  not 
wish  to  make  too  much  of  such  a  contention,  though  I  shall 
bring  up  the  point  again  presently.  All  I  want  to  do  just 
here  is  to  make  still  clearer  the  meaning  of  my  view  that 
James  was  organismal  in  spirit,  though  not  wholly  so  in  for- 
mal statement.  To  me  one  of  the  strongest  evidences  of 
this  was  his  obvious  effort,  as  indicated  by  these  and  many 
other  passages  in  many  other  writings  than  his  Psi/choJocj?/, 
to  describe  fully  the  phenomena  with  which  he  chanced  to 
deal.  As  I  have  remarked  in  substance  so  many  times  in  this 
book,  one  of  the  most  unmistakable  signs  of  the  elementalist 
attitude  in  biologv  is  incomplete  and  more  or  less  perverted 


322  The  Unity  of  the  Organism 

description.  And  nowhere,  perhaps,  in  the  whole  biological 
realm  is  there  a  better  chance  for  description  of  the  genuine- 
ly natural  history,  organismal  kind — the  kind  a  cardinal 
motto  of  which  is  "neglect  nothing,"  than  in  this  very  field 
of  human  emotions,  especially  of  one's  own  emotions.  Nor 
can  I  refrain  from  reminding  the  reader  that  one  of  the 
master  works  in  this  field  is  Darwin's  The  Expression  of  the 
Emotions  in  Man  and  Animuls,"^^  and  that  while  a  leading 
motive  of  its  author  was  to  interpret  the  emotions  in  ac- 
cordance with  the  theory  of  descent  and  the  natural  selec- 
tion hypothesis,  probably  the  most  lasting  value  of  the  work 
is  from  its  fullness  and  excellence  as  a  natural  history  de- 
scription of  the  emotions  and  their  objective  expression. 

As  to  the  fact  of  vital  interdependence  between  psychic 
life  and  physical  life  through  the  emotions,  personal  experi- 
ence and  observation,  backed  up  and  supplemented  by  many 
authoritative  writings,  among  which  those  of  Darwin  and 
James  stand  out  strongly,  there  seems  no  longer  any  room 
for  question.  The  role  of  the  emotions  as  between  "Body" 
and  "Soul"  may  be  crudely  likened  to  the  splice  which  a  skill- 
ful sailor  weaves  into  two  pieces  of  rope  in  joining  them  so 
that  there  shall  be  no  knot  and  as  great  strength  as  in  any 
other  part  of  the  rope.  In  the  recent  period  of  psychology 
■ — of  so-called  physiological  psychology — we  have  frequently 
heard  about  psychology  "without  a  Soul ;"  and  such  an  idea 
has  seemed  repugnant  to  many  persons.  But  if  we  could 
show  that  this  modern  psychology  is  "without  a  Body"  by 
the  same  token  that  it  is  "without  a  Soul,"  the  legitimate  mis- 
givings about  the  soullessness  of  the  psychology  ought  to  be 
allayed.  And  really  the  organismal  conception  of  psychic 
life  is  seen,  especially  when  we  examine  it  in  the  phase  of  the 
emotions,  to  amount  to  such  a  composition  of  the  Body-Soul 
antithesis.  "Body"  we  can  see,  as  it  figured  in  the  old  psy- 
chology, virtually  signified  what  we  usually  mean  by  corpse, 
or  cadaver.     "The  Body,"  in  that  sense  was  not  alive  at  all. 


Sketch  of  an  OrganismdJ  Throri/  of  Consciousui'ss     323 

It  was  not  alive  because  all  the  life  was  taken  out   of  it    (  hy 
the  theoretical  antithesis)  and  put  into  ''The  Soul." 

Glance  at  the  Equilihrative  Interaction  Between  'Ihtdif'  (iiid 

'\Sour 

Going  forward  from  sucli  predominantly  observational 
descriptions  of  psychic  life  in  its  emotional  phase  as  those  of 
Darwin  and  James,  to  such  experimental  descriptions  as 
those  being  produced  by  the  investigations  of  Pawlow,  of 
Crile,  and  especially  of  Cannon,  we  are  getting  considerable 
insight  into  the  rationale  of  how  "Body"  and  ''Soul"  vitalize 
each  other.  Modern  researches  on  the  physiology  or  the 
psychology  (which  one  calls  it  depends  entirely  on  the  direc- 
tion of  his  approach)  of  psychic  life  is  revealing  something 
of  the  why  and  how  of  the  poet's  instinctive  perception,  "Soul 
needs  Body  as  much  as  Body  needs  Soul."  Only  one  aspect 
of  this  "why  and  how"  need  be  noticed  in  the  present  discus- 
sion. That  is  the  fact  of  the  balancing  off'  of  antagonistic 
emotions  to  make  the  normal  emotional  life  just  as  reflex- 
actions  and  instinctive  actions  are  largely  phenomena  of 
equilibration,  or  balancing-ofF. 

It  should  be  recalled  that  we  have  found  this  antagonistic- 
equilibrative  principle  to  run  through  the  entire  neuro-psy- 
chic  life.  In  the  strictly  reflex  phase  the  mode  of  operation 
of  the  opposing  muscles,  the  flexors  and  extensors  of  the 
limbs,  as  brought  out  by  Sherrington,  was  cited  as  a  good 
illustration  of  the  princi])le.  A  manifestation  of  the  ])rin- 
ciple  in  a  broader  way,  as  measured  by  the  extent  of  organic 
parts  involved,  was  seen  in  the  relation  of  the  vagal  (cranial) 
and  splanchnic  (thoracico-lumbar )  autonomies,  as  empha- 
sized by  Cannon  (Chap.  19,  this  book)  this  illustration  being 
chiefly  in  the  reflex  phase.  In  a  yet  higher  })hase  we  saw, 
again  from  Cannon's  work,  the  })rineiple  in  o]H'ration 
through  the  emotions  (Chap.  TS)  thus  bringing  it  up  to  the 


324  The  Unity  of  the  Organism 

phase  of  lower  conscious  life. 

The  reader  should  not  forget  the  insistence  throughout 
our  presentation  of  these  antagonistic  phenomena,  that  al- 
ways the  oppositions  and  antagonisms  and  competitions  are 
fundamentally  constitutive  as  to  the  normal  organism.  Even 
the  most  pronounced  of  them  are  yet  in  the  interest  of  the 
organism  as  a  whole.  They  are  always  partial  phenomena 
relative  to  the  whole  organism.  They  have  evolved  in  strict 
accordance  with  and  sub-ordination  to  the  fundamental  na- 
ture of  the  organism  in  its  totality.  The  opposing  muscles 
of  our  limbs  can  not  break  or  tear  one  another  under  normal 
conditions.  Even  antagonisms  among  the  parts  of  the  or- 
ganism are  possible  because  the  parts  belong  to  the  organism. 
The  antagonisms  of  the  parts  do  not  produce  the  organism, 
primarih^,  but  are  themselves  produced  by  the  organism,  or 
at  least,  are  a  portion  of  the  means  or  methods  by  which  the 
organism  lives  and  enlarges,  develops  and  functions.  All  this, 
be  it  noticed,  holds  not  merely  as  touching  purely  physical 
organization  *  but  as  to  the  entire  gamut  of  psychic  life, 
at  least  up  to  and  including  instinctive  and  emotional  life. 

Support  of  the  Hypothesis  by  the  Physico-Chemical  Con- 
ception of   the   Organism 

This  prepares  us  for  the  final  step  of  switching  the  discus- 
sion from  the  psycho-conscious  aspect  of  life  to  the  bio- 
physico-chemical  aspect.  The  place  in  our  discussion  to 
which  this  return  naturally  takes  us  is  that  wherein  we  con- 
sidered the  organism's  chemical  nature  as  interpreted  by  phy- 
sical chemistry.  That  interpretation  has  been  presented  by 
several  physiologists  but  with  special  insight  and  cogency  by 
F.  G.  Hopkins.  For  example,  our  citation  in  Chapter  4  of 
the   statement   that   the   conception    of   the   organism   as   a 

*  Recall  the  discussions  of  growth  and  chemico-functional  integration, 
chapters  17,  18,  and  19. 


Sketch  of  an  Orgauismal  TJicory  of  Consciousness     iVl') 

cliemical  laboratory  "is  rajjidly   «raininfr  ground,''  should  \yc 
recalled,  as  should  also  the  o])iiiion  of  lloi)kins:     "the  chem- 
ical response  of  the  tissues  to  the  chemical  stimulus  of  foreign 
substances  of  sim])le  constitution  is  of  ])r()fouiid  biological 
significance,"  and  tliat  further  study  of  the  phenomena  "must 
throw  vivid  light  on   the  potentialities  of  the  tissue  labora- 
tories."^^    So  far  this  chemical  laboratory  conception  of  the 
tissues  may  be  said  to  be  strictly  chemical ;  but  let  us  recall 
what  the  interpretation  is  when  it  passes  from  chemistry  in 
the  exclusive  sense  to  physical  chemistry  and  becomes  more 
specific  as   to  the  laboratory   apparatus,   as   one  may   say, 
through  which  the  "tissues"  work.     In  other  words,  recall 
the   conception   of  the   cell   and   its   mode   of   operating,   as 
viewed    by    physical    chemistry.      The    quotations    given    in 
Chapter     4  may  well  be  repeated  in  part:  ".   .   .   the  living 
cell  as  we  now  know  it  is  not  a  mass  of  matter  composed  of  a 
congregation   of  like  molecules,   but   a  highly   differentiated 
system ;  the  cell  in  the  modern  phraseology  of  physical  chem- 
istry, is  a  system  of  coexisting  phases  of  different  consti- 
tutions." ^^     Then  from  this  review  our  own  contention,  set 
forth  especially  in  Chapter  7,  that  wherever  in  such  state- 
ments as  those  just  quoted  from  Hopkins  "the  term  ceU  oc- 
curs the  term  organism  really  ought  to  be  used." 

It  is  important  for  our  cause  generally  that  the  full 
weight  of  our  argument  in  support  of  the  view  that  on  the 
strictly  physical  plane,  the  organism  rather  than  the  cell 
is  really  the  equilibration  system  toward  which  physico- 
chemical  knowledge  is  tending,  should  be  In  liie  reader's  con- 
sciousness. At  this  point  if,  consequently,  this  Is  not  so, 
he  is  urged  to  read  what  Is  said  on  tlie  ])oint  in  Chapters 
4  and  7  especially. 

Our  central  purpose  now  is  to  show  that  tlie  organismal 
hypothesis  of  consciousness  articulates  directly  and  natur- 
ally with  the  same  conception  of  the  organism,  rndoubtedly 
it  is  in  the  emotional  phase  of  psychic  life  that  this  articu- 


^^6  The  Unity  of  the  Organism 

lation  is  most  open  to  common  obsers^ation.  Compare,  for 
example,  James'  "Our  whole  cubic  capacity  is  sensibly  alive; 
and  each  morsel  of  it  contributes  its  pulsations  of  feeling, 
dim  or  sharp,  pleasant,  painful,  or  dubious,  to  that  sense  of 
personality  that  every  one  of  us  unfamiliarly  carries  with 
him,"  with  Hopkins'  "On  ultimate  analysis  we  can  scarcely 
speak  at  all  of  living  matter  in  the  cell ;  at  any  rate,  we 
cannot,  without  gross  misuse  of  terms,  speak  of  the  cell- 
life  as  being  associated  with  any  one  particular  type  of  mole- 
cule. Its  life  is  the  expression  of  a  particular  dynamic  equil- 
ibrium which  obtains  in  a  polyphasic  system  .  .  .  'life'  as  we 
instinctively  define  it,  is  a  property  of  the  cell  as  a  whole, 
because  it  depends  upon  the  organization  of  processes,  upon 
the  equilibrium  displayed  by  the  totality  of  the  coexisting 
phases."  ^^  Also  compare  Hopkins'  statement  that  among 
the  different  "phases"  of  the  cell  in  which  its  life  inheres, 
"are  to  be  reckoned  not  only  the  differentiated  parts  of  the 
bio-plasm  strictly  defined  (if  we  can  define  it  strictly),  the 
macro-and-micro-nuclei,  nerve  fibers,  muscle  fibers,  etc.,  but 
the  materials  which  support  the  cell  structure,  and  which 
have  been  termed  metaplastic  constituents  of  the  cell,"  with 
James'  "each  morsel"  of  our  cubic  capacity  "contributes  its 
pulsations  of  feeling,  etc." 

The  congruity  of  these  statements  is  apparent  even  when 
taken  as  here  exhibited ;  that  is,  each  as  standing  by  itself 
at  about  the  two  extremes  of  the  scale  of  life.  When,  how- 
ever, they  are  viewed  in  connection  with  my  general  argument 
that  "cell"  in  Hopkins'  statement  ought  to  be  replaced  by 
"organism" ;  and  in  connection  with  what  we  have  learned 
from  Cannon  and  others  about  the  mechanism  by  means  of 
which  the  organism  operates  in  the  phase  of  conscious  emo- 
tion, it  seems  as  though  our  organismal  hypothesis  of  con- 
sciousness comes  near  to  a  demonstration.  And  so  far  as 
ordinary  descriptive  natural  history  is  concerned,  I  believe 
this  to  be  true.     However,  I  recognize,  keenly  enough,  that 


Sketch   of  an  Orf/auismnJ  Throrif  of  Con.^rinn.siu.ss      .'527 

from  the  standpoint  of  hio-clicniistrv,  and  plivsiolocry,  }uid 
also  from  that  of  ])hih)s()j)hv  In  Ww  traditional  sense,  tliat 
demonstration  is  not  only  far  away,  hut  is  attainahlc,  if  at 
all,  only  by  surmoiintinn-  \cvy  formidahlc  difficult  its.  So  I 
reassure  the  dubious  reader  that  all  I  am  elaimimr  is  that 
my  two  propositions  a})()ut  the  nature  of  consciousness  to- 
gether constitute  a  legitimate  scientific  liypothesis. 

Personality  and  Elevientary  Chemical  Substances 

With  both  the  physico-chemical  aspect  and  the  psychical 
aspect  of  our  hypothesis  now  before  us  more  fully  and 
sharply  than  they  have  been  hitherto  we  will  examine  an  ob- 
jection to  it  which  I  apprehend  will  be  the  most  serious  the 
hypothesis  will  meet;  namely'  that  to  the  proposition  that 
each  individual  organism  has  the  value  in  a  chemical  sense 
of  an  elementary  substance.  And  since  this  objection  will 
probably  be  more  intolerant  and  stubborn  from  the  side  of 
physics  and  chemistry  than  from  that  of  natural  history  and 
psychology  I  will  adjust  my  remarks  with  reference  to  the 
opposition  as  thus  anticipated. 

The  considerations  I  am  going  to  present  might  have  been, 
in  strict  expository  coherence,  presented  as  a  part  of  my 
discussion  of  the  uniqueness  of  the  individual  consciousness 
as  marked  by  its  necessary  privacy  and  its  difference  from 
all  other  individual  consciousness.  What  we  are  now  to 
emphasize  is  the  fundamentality  of  objective  as  contrastetl 
■with  subjective  personality  of  sucli  highly  develo])ed  animals 
as  song  birds,  domesticable  animals,  and  civilized  man. 

A  complete  definition  of  ''personality'"  is  not  obligatory 
for  our  purpose.  Only  this  much  need  be  said  about  the 
meaning  we  shall  give  the  word:  First,  we  deny  the  right 
claimed  by  some  authors  to  make  personality  purely  j)sy- 
chical,  or  spiritual — a  thing  of  the  "inner,"  or  '*dee])er'* 
self;  "Self"  that  is,  in  a  thorough-going  subjectivistic  sense. 


328  The  Unity  of  the  Organism 

It  is  on  this  ground,  as  I  understand,  that  some  psycholo- 
gists, as  G.  F.  Stout,  and  apparently  C.  Lloyd  Morgan,^^ 
deny  personality  to  animals.  All  I  will  say  on  this  question 
here  is  that  I  am  quite  sure  that  every  close  observer  of  the 
higher  animals  will  recognize  that  if  he  undertakes  to  give  a 
truly  full  report  of  his  observations  on  their  behavior  he  will 
have  to  speak  of  the  personality  of  some  at  least  of  them 
just  as  he  would  of  the  personality  of  observed  human  beings, 
or  he  will  be  obliged  to  call  the  same  thing  by  some  other 
name — a  kind  of  procedure  against  which  we  have  spoken 
strongly  throughout  this  volume.  For  us,  whatever  person- 
ality may  be,  we  must  conceive  it  to  be  founded  upon,  and 
conformable  to,  the  organism.  "Organism"  must  be  the  more 
inclusive  term.  "Person"  must  stand  to  "Organism"  in  the 
logical  relation  of  species  to  genus. 

Another  meaning  of  personality  in  this  particular  dis- 
cussion will  concern  the  uniqueness  of  each  organism  as  to  its 
psychical  attributes  regarded  in  their  totality.  By  unique- 
ness I  mean  not  merely  the  fact  that  each  organism  is  itself, 
perceptually  regarded,  but  that  it  is  not  a  replica,  a  dupli- 
cate of  any  other.  It  is  not  only  another  organism  but  it  is 
in  some  measure  a  different  other  organism.  For  the  benefit 
of  those  physical-  and  metaphysical-minded  readers  who  have 
never  informed  themselves  much  about  the  facts  of  natural 
history  and  have  never  tried  seriously  to  think  in  the  nat- 
ural history  manner  I  would  remark  that  what  I  have  just 
said  concerning  the  uniqueness  of  the  individual  organism 
is  only  re-asserting  in  a  more  refined  way  what  botany  and 
zoology  have  recognized  more  or  less  definitely  since  Dar- 
win's time  at  least,  and  have  partially  expressed  in  the  terms 
"individual  difference"  and  "individual  variation." 

With  this  we  come  to  the  cardinal  point:  If  individual 
animal  organisms,  especially  individual  humans  under  civi- 
lization, he  contemplated  with  due  heed  to  the  motto  ^'neglect 
nothijig''  the  conviction  will  be  reached  that  each  and  every 


Sketch  of  an  Organismal  Thconj  of  Conscious luss     329 

one  has  literally  as  much  of  uniqueness  about  it  as  has  an 
elementary  chemical  substance. 

In  order  to  bring  out  the  trutli  of  this  statement  we  must 
exhibit,  in  the  regular  natural  history  manner,  the  resem- 
blances and  differences  between  cliemical  elements  on  tlie  one 
hand  and  the  resemblances  and  differences  between  human 
beings  on  the  other,  and  tlien  pool  the  results  of  these  com- 
parisons. 

To  the  carrying  out  of  this  enterprise  the  so-called  peri- 
odic law   in   chemistry   is   of  very   great   importance.      The 
essence  of  this  law,  stated  from  the  natural  history  stand- 
point, is   that   the   chemical  elements   range   themselves   into 
natural  species  and  genera  after  much  the  fashion  that  plants 
and  animals  do ;  and  that  the  classification  is  based  mostly  on 
the  chemical  attributes  of  the  substances,  but  partly  on  their 
physical  attributes  also.     Thus  the  "halogen  group,"  that  to 
which  lithium,  sodium,  and  potassium  belong,  is  a  genus  in 
the  sense  of  descriptive  natural  history,  its  species  being  the 
substances  mentioned  with  others  not  enumerated.     Also  the 
group  often  spoken  of  in  chemical  laboratories  as  "the  iron 
group" — the  genus  containing  the  species  iron,  cobalt,  nickel, 
platinum,  etc.,  illustrates  the  point.    Two  species  of  the  last 
genus,  iron  and  nickel,  will  be  used  in  our  stud3\     Let  us 
compare  some  household  utensil  made  of  iron  with  a  similar 
one  made  of  nickel.     For  the  ordinary  uses  to  which  these 
implements   would   be    put    the   difference   between    the   sub- 
stances   of  which  they   are  made   would   hardly   Ix^   noticed. 
The  higher  specific  gravity  of  nickel  (8.5  plus)   is  so  sHght 
as  compared  with  that  of  iron  (7.8)  that  the  greater  welglit 
of  the  nickel  im])lement  would  ])rol)ably  not  1h^  noticed.     Nor 
would  the  slightly  lower  melting  j)oint  of  nickel  nor  its  much 
lower   magnetic    capacity    be    recognized.      The   most    avad- 
able  distinguishing  difference  is  in  color,  the  ordinary  house- 
keeper answering  you,  if  you  ask  how  she  knows  a   nickel 
from  an  iron  implement,  that  the  nickel  piece  is  silvery  bright 


330  The  Unity  of  the  Organism 

while  the  iron  piece  is  black. 

See  now  what  this  means.  Actually,  as  is  well  known  to 
every  beginning  student  in  analytical  chemistry,  these  two 
metals  are  very  similar  in  color  as  well  as  in  other  physical 
attributes — so  much  so,  in  fact,  that  some  authors  apply 
the  same  term  "silver  white"  to  both.  What  a  housekeeper 
really  means  when  she  says  she  knows  one  implement  to  be 
of  nickel  because  it  is  bright  and  the  other  to  be  of  iron 
because  it  is  black,  is  that  she  is  depending  on  a  chemical 
rather  than  a  physical  attribute  for  a  distinguishing  mark ; 
the  attribute,  that  is,  in  virtue  of  which  iron  is  acted  upon 
much  more  readily  by  oxygen  in  the  presence  of  moisture 
than  is  nickel.  The  much  greater  liability  of  iron  than 
nickel  to  tarnish  and  rust  is  a  chemical  rather  than  a  phy- 
sical difference  between  them.  This  fact,  namely  that  of 
the  dependence  of  distinguishing  differences  between  sub- 
stances more  upon  chemical  than  upon  physical  attributes  is 
of  very  wide  applicability  in  nature,  and  is  greatly  impor- 
tant both  scientifically  and  philosophically. 

Now  turn  from  comparing  these  two  elementary  chemical 
substances  to  a  comparison  of  any  two  human  organisms,  or 
persons  who  might  be  members  of  a  household  to  Avhich 
the  implements  might  belong.  And  make  the  comparison  first 
on  the  basis  of  the  ph3^sical  attributes  just  as  we  began 
comparing  the  implements  of  nickel  and  iron.  Does  any 
reader  doubt  that  he  would  find  it  much  easier  to  distinguish 
the  persons  than  the  metals.'^  As  to  purely  morphological, 
that  is,  physical  differences  between  almost  any  two  persons 
(with  the  possible  exception  of  certain  rare  instances  of 
"identical"  twins),  there  is  no  room  for  question.  General 
shape  of  head,  face  and  features,  and  the  size  and  propor- 
tions of  the  various  parts  of  the  body  furnish  many  unmis- 
takable distinguishing  attributes. 


Sketch   of  an  Orcfduisnud  'llicorij  of  Consciousness     3551 

On  the  Psychology  of  Subjective  ami  Objective  Personality 

But  unerring-  as  arc  the  (lifrcrcntiatin^  marks  on  tlie 
physical  side,  such  marks  arc  few  as  compared  with  those 
on  the  psychical  side.  Noting  first  certain  merely  physico- 
psychical  differences  think  of  the  manners  of  speech  and  of 
hand  writing,  to  mention  only  two  items!  I'ndoubtedly  these 
differences  arc  to  a  considerable  extent  ])hysical  hut  no  one 
would  seriously  question  that  psycliical  factors  come  in  all 
along  the  line.  This  is  perha])s  most  obvious  in  sj)eech  as 
evidenced  by  voice  modulations,  intonations,  gesticulations, 
and  facial  and  bodily  expressions.  Again,  differentials  are 
everywhere  recognizable  in  responses  to  sensory  stimuli, 
especially  in  the  matter  of  reaction-time.  There  are  the 
quick  and  accurate  persons,  and  the  quick  and  inaccurate 
ones ;  and  there  are  the  slow  and  accurate  and  the  slow  and 
inaccurate  types,  to  go  only  a  step  in  descrii)ti()n  and  classi- 
fication on  this  basis. 

Then  we  proceed  to  compare  the  unequivocal  psychical 
phases  of  life:  the  feeling,  the  emotional,  the  esthetic,  the 
religious,  and  the  intellectual  phases.  Here  we  pass  into 
a  realm  of  what  might  properly  be  called  objective  i)rivacy 
in  psychology,  individuals  for  the  study  of  which  would  be 
largely  the  student's  most  intimate  and  most  enduring  friends 
and  associates,  human  and  animal.  Such  a  psyciiology  would 
be  undeniably  so  particular  and  intimate  that  much  of  it 
would  be  unpublishable  even  if  it  had  an  interest  beyond 
the  few  persons  concerned.  At  the  same  time  there  are 
portions  of  it  of  great  jiublic  importance,  one  such  ])() 
tion  being  exactly  what  we  are  in  need  of  in  tlu-  present  dis- 
cussion. I  refer  to  the  exceedingly  familiar  hut  sciintifically 
much  neglected  definite  and  sustaim«d  psycliical  differences 
of  individuals  who  by  reason  of  Iniiig  in.  iiilurs  of  tlu'  s.imo 
household  or  same  small  community  are  subject  to  nearly 
identical  influence  so  far  as  concerns  such   fundamental  en- 


some 
r- 


^^^  The  Unity  of  the  Organism 

vironic  factors  as  food  in  the  narrow  sense,  drink,  air,  light 
and  temperature.  The  duty  before  us  is  that  of  testifying  to, 
of  viseing,  the  objectively  psychical  individual  as  we  did  the 
subjectively  psychical  individual  earlier  in  this  sketch. 
"What  is  needed,"  writes  Sellars,  "is  not  vague  statements  to 
the  effect  that  individuals  cannot  be  separated  or  that  they 
are  aspects  of  one  another,  but  definitions  and  analyses."  ^'^ 
Sellars  is  here  raising  his  voice  against  the  tendency  in 
present-day  social  psychology  to  make  the  individual  a  kind 
of  incident  in  the  social  order,  a  by-product  of  Society.  It 
is  a  satisfaction  that  the  regular  course  of  my  psychological 
argument  has  brought  me  to  where  I  also  may  contribute 
something  to  the  definition  and  analyses  essential  to  check- 
ing the  tendency  indicated  by  Sellars.  If  it  can  be  shown 
biologically  and  psychologically  all  in  one  that  personality 
is  indubitably  objective,  both  substantively  and  kinetically, 
not  only  the  Individual  but  Society  will  be  the  gainer,  I  am 
very  sure.  For  my  contribution  we  will  examine  in  outline 
what  may  appropriately  be  called  the  action-system  (adopt- 
ing and  expanding  Jennings'  term)  as  it  manifests  itself  in 
a  small  homogeneous  group  of  human  beings.  Our  study  will 
be,  in  other  words,  one  in  domestic  and  neighborhood  psy- 
chology. 

The  "material"  in  this  instance  must  be  my  own  household 
and  the  handful  of  persons  constituting  the  colony  of  the 
Scripps  Institution  for  Biological  Research.  This  group 
is  rather  specially  favorable  for  such  a  study  in  that  its 
geographic  severance  from  other  groups,  and  its  strictly 
rural  habitat  give  it  an  exceptionally  natural,  simple,  and 
uniform  environment.  The  analysis  might  run  along  any 
one  or  all  of  several  axes;  but  our  purpose  will  be  accom- 
plished by  following  one  only.  That  one  shall  be  the  reac- 
tion, the  behavior,  of  individual  members  of  the  group  in 
response  to  the  stimulus  of  the  world  war.  Were  complete- 
ness to  be  aimed  at  in  the  analysis,  every  individual  in  the 


Sketch  of  an  Organismal  Theory  of  Comcwnsnesi     333 

group  would  have  to  be  considered.  Sueli  a  treatment  would 
be  highly  instructive  but  s})ace  limitations  forbid  us  going  to 
such  length.  We  must  restrict  ourselves  to  a  few  of  the 
more  pronouncedly  individualistic  behaviors  and  must  treat 
even  these  in  a  very  sketchy  fashion.  To  Ik'  remarked  at 
the  outset  is  the  fact  that  every  nicnilHr  of  the  group  is 
deeply  loyal  to  America  and  to  the  cause  of  the  Allies.  On 
the  very  door-sill  of  the  examination  we  recognize  two  well- 
differentiated  aspects  to  each  i)erson's  action-sy.^tem,  namely 
an  aspect  of  connnonality  for  nearly  all  members  of  the 
group ;  and  an  aspect  of  very  pronounced  differentiality  for 
many  of  them. 

Behaviors-in-common  will  receive  attention  first.  In  the 
uniform  growth,  from  the  very  beginning  of  the  struggle  in 
August,  1914,  of  belief  in  the  general  rightness  of  the  cause 
of  the  Entente;  of  realization  of  the  meaning  of  the  strufrtrlc; 
and  of  sentiments  and  resolutions  of  devotion  to  the  foreimi 
nations  with  which  our  nation  is  finally  joined,  these  experi- 
ences have  been  very  much  at  one.  To  be  sure  this  connnon- 
ness  has  fallen  far  short  of  identitv.  But  as  to  essentials 
resemblance  has  been  far  greater  than  difference.  For  ex- 
ample every  adult  has  accepted  unhesitatingly  his  and  her 
obligations  to  the  Red  Cross;  to  the  appeals  for  aid  from 
Belgium,  France,  and  the  other  despoiled  countries;  to  the 
increasing  cost  of  living;  to  the  buying  of  Government 
Bonds ;  and  to  the  appeals  and  regulations  of  the  Food  Ad- 
ministration. Naturally  there  has  been  difference  in  the 
particular  way  and  extent  of  response  of  each  in  these  mat- 
ters;  but  in  essence  there  has  been  nothing  differential. 

We  turn  now  to  behavior-not-in-connnon  :  behavior,  that 
is,  which  has  differentiated  the  members  i)ersonally  with 
great  sharpness.  This  examination  is  mucli  more  important 
for  the  subject  in  hand.  The  reference  here  is  to  each  one's 
"bit"  as  the  common  phrase  had  it  when  our  country  was 
first  enterinjr  the  conflict.     The  "war  work"  (as  the  exprcs- 


334  ■    The  Unity  of  the  Organism 

sion  has  gradually  become  with  the  advance  toward  the  cli- 
max of  the  gigantic  struggle)  into  which  each  has  gravitated 
has  much  the  appearance  of  the  naturalness  and  inevitability 
presented  by  the  falling  of  a  stone  or  the  flowing  of  water. 
The  case  grows  so  significant  at  this  point  that  I  must  par- 
ticularize somewhat  more  than  I  have  heretofore.    A  becomes 
an   acknowledged  leader  in  "drives"   for  Red   Cross   funds, 
Liberty  Bond  sales,  etc.     B  becomes  a  regular  consultant 
on  the  knitting  of  Red  Cross  articles.     C  is  a  highly  skilled 
deviser  and  maker   of  dishes   from   "substitute"   foods.      D 
is  appointed  an  official  of  the  National  Food  Administra- 
tion.    E  becomes  an  official  teacher  of  girls  and  women  as  to 
the  peculiar  duties  and  obligations  of  their  sex  in  war  times. 
F  concentrates  nearly  the  whole  of  his  physical  energy  upon 
an  elaboration  of  the  view  that  a  victory  over  Germany  and 
her  allies  cannot  be  really  complete  without  being  spiritual 
as  well  as  material — that  the  philosophy  or  theory  of  life 
being  fought  for  by  Germany  must  be  overthrown  as  well  as 
her  armed  forces.     Of  the  forty  adult  members  of  the  group 
fully  one-half  have  been  incited  in  a  special  degree  to  some 
activity  that  has  a  distinct  personal  character,  some  of  these, 
as  above  indicated,  being  very  pronouncedly  so.     The  per- 
sonality of  these  reactions  comes  to  view  most  distinctly  in 
the  fact,  absolutely  certain  to  an  observer  whose  acquain- 
tance with  the  persons  has  been  intimate  and  has  extended 
over  some  years,  that  no  one  of  those  who  has  settled  into 
one  of  the  special,  definite,  and  important  pieces   of  work 
could  wholly  replace  any  of  the  others  in  their  special  tasks. 
Probably  each  could  do  something  at  the  "job"  of  any  of 
the  others  were  conditions  such  as  to  force  him  to  try;  but 
success  under  such  conditions  would  surely  be  partial,  very 
much  so  in  some  of  the  cases. 

This  automatic  definition  and  classification  of  persons  sub- 
ject to  a  common  major  stimulus,  with  nearly  the  same  gen- 
eral environic  conditions,  and  with  almost  complete  freedom 


Sketch  of  an  Orgatilsrual  Thcor/j  of  Consciousness     335 

of  action  so  far  as  concerns  tlic  particular  stimulus,  sccnis 
to  me  a  phenomenon  of  very  great  imi)ortance  since  it  de- 
pends upon  princij)les  of  organic  beings,  especially  upon 
principles  of  civilized  nian's  "being,"  which  are  well-nigh  if 
not  entirely  universal,  I  am  sure.  Undoubtedly  the  phenom- 
enon is  often  much  obscured  through  counteracting  ele- 
ments in  the  environment,  especially  in  social  customs,  eco- 
nomic conditions  and  general  education  among  civilized  men. 
But  in  spite  of  all  these,  attentive  observation  will  nearly 
always  be  able  to  recognize  it.  Highly  significant  is  it  as 
bearing  on  this  particular  aspect  of  the  matter,  that  the 
niches  finally  found  by  most  of  the  persons  were  obviouslv 
determined  to  some  extent  by  long  continued  previous  activi- 
ties and  unmistakable  natural  "gifts." 

Another  noteworthy  fact  is  the  clear  indication  of  not 
mere  acceptance,  but  positive  satisfaction  on  the  part  of 
most  if  not  all  the  persons,  once  they  are  "settled"  to  their 
"jobs,"  this  satisfaction  prevailing  despite  the  strenuousness, 
perplexity,  and  wear-and-tear  entailed.  During  the  first 
weeks  of  America's  plunge  into  the  maelstrom  the  anxious 
psychical  casting  about  in  our  little  group,  as  throughout 
the  whole  land,  presents  to  the  anthropological  biologist  as 
he  looks  back  upon  it  a  case  of  trial  and  error  on  a  gigantic 
scale,  the  scene  being  replete  with  jumbled  elements  of  noble 
zeal,  splendid  efficiency,  mis-expenditure  of  strength  and 
funds,  and  ludicrous  proposals.  But  out  of  this,  as  out  of 
this  unprecedented  instance  of  world-wide  "struggle  for  ex- 
istence," there  is  quite  sure  to  come,  indeed  is  coming,  as  one 
of  its  first  fruits,  personalit/j  more  real  and  powerful  and 
fuller  of  grandeur  than  ever. 

While   personalities    come    forth   with   special    distinctness 
of  outline  and  forcefulness  of  ex])ression  during  occasional 
events   of   vast    import    to    the    race    like    tiu'    |)rcs(Mit    war    m 
volving  literally    the   whole   civilized    ])()rtion    of    the    human 
species,  yet  I   would   insist   that   the   ditrerence   Ix^tween   the 


^^6  The  Unity  of  the  Organism 

manifestations  at  such  times  and  at  ordinary  times  is  al- 
most entirely  one  of  degree,  rather  than  of  essential  nature. 
The  attentive  observer  will  not  fail  to  find  personalities  as 
here  understood  always  and  everywhere,  no  matter  how  sim- 
ple and  lowly  the  lives,  and  monochrome  the  external  condi- 
tions. In  little  details  of  intelligent,  but  still  more  of  reflex, 
instinctive,  and  emotional  life,  all  of  which  compounded  to- 
gether makes  what  we  often  call  temperament,  the  keen  and 
sympathetic  observer  will  always  see  persons  in  the  deep 
sense  here  indicated.  Not  the  transcendent  genuises  merely, 
the  Aristotles,  the  Shakespeares,  the  Napoleons,  have  the 
right  to  be  called  personalities,  because  of  the  unique  powers 
with  which  they  are  endowed ;  but  each  and  every  one  of  civi- 
lization's humblest-ranked  myriads,  and  each  and  every 
nature-tutored  denizen  of  the  virgin  forest,  of  the  untilled 
plain,  and  of  the  unregenerate  desert,  have  the  same  right- 
in-kind. 

Personality  and  the  '^Breath  of  Life""  Viewed  in  the  Light 
of  Physical  Chemistry  of  the  Organism 

Swinging  the  discussion  back  now  on  the  physico-chemical 
aspect  of  the  organism,  I  recall  first  the  truth  alluded  to  a 
little  while  ago,  namely,  that  it  is  preeminently  the  chemical 
rather  the  physical  attributes  of  elementary  inorganic  sub- 
stances which  furnish  the  distinguishing  marks  of  these  sub- 
stances. Even  in  the  inorganic  world  we  saw  that  substances 
are  most  readily  and  decisively  differentiated  from  one  an- 
other by  the  transformation-products  resulting  from  the 
reaction  of  the  substances  upon  one  another.  "Transforma- 
tion of  energy,"  using  a  form  of  expression  favored  by  the 
disembodying  tendencies  in  recent  chemical  theory,  is  the 
most  distinctive  thing  about  all  chemistry,  inorganic  as  well 
as  organic.  The  oxidation  and  other  chemically  reactive 
changes  and  products  of  nickel  and  iron,  we  noticed,  are  the 


Sketch  of  an  Orgamsmal  Theory  of  Couscwusiu\ss     3S7 

most  difFerentiativc  things  about  these  metals.  Let  us  push 
the  application  of  tliis  criterion  of  difference  a  little  farther 
in  comparing  human  persons.  We  give  energy-transforma- 
tion and  work  performed  a  leading  place  here  also.  And 
lx?ing  naturalistically  chemical  rather  than  chemically  chem- 
ical we  are  forced  to  touch  the  ''high  spots"  only  at  first 
regardless  of  what  may  be  in  between  them.  We  are  free  to 
seize  upon  the  end  or  completed  products  of  the  reactions 
and  transformations.  What  reaction-products,  I  ask,  of 
nickel  and  iron  towards  any  other  substance  or  set  of  condi- 
tions are  more  unlike  than  the  reaction-products  of  an  effi- 
cient Department-of-Justice  official,  let  us  say  and  an  ef- 
ficient food  conserving  house-keeper,  in  this  time  of  common 
national  danger .'^  Yet  these  diverse  products  may  come  from 
not  only  the  same  danger  stimulus,  but  likewise  from  as  nearly 
identical  physico-chemical  environic  stimuli  as  it  is  possible 
to  secure.  Were  official  and  house-keeper  to  eat  of  the  sam<' 
food,  drink  of  the  same  fluids,  breathe  of  the  same  air,  and 
be  subject  to  the  same  temperatures  month  in  and  month 
out  the  difference  in  product  would  not  be  a  whit  less. 

So  stands  the  case  when  viewed  in  its  "high  places"  only. 
But  the  high  places  are  as  real  places  as  any  whatever.  No 
realities,  it  matters  not  how  obscure  or  subtle,  pertaining  to 
the  intermediate  places,  can  make  the  high  places  other 
than  what  thej'-  are.  Judging  human  beings  by  what  they  do, 
by  work  done  through  the  transformation  of  the  substances 
and  energies  which  they  take  from  the  external  world,  their 
personalities  *»\'e  surely  not  less  well-attested  than  are  the 
individualities  v,r  elementary  chemical  substances.*  Hut  it 
will  not  do  to  be  satisfied  with  touching  the  high  placis  in 
this  rather  jaunty  fashion.     Some  attention  must  be  given  to 

*  A  rather  full  discussion  of  tlie  point  here  tonclied  may  l)e  f«iund  in 
my  essay,  The  Jfiffher  Uscfiiliic'^ti  of  Srinur.  where  I  rai.se  anil  try  to 
answer  the  query,  "What  is  nature  because  man  is  a  part  of  it?"  Per- 
haps a  less  ambiguous  way  of  asking  the  question  would  be,  "What  must 
nature  be  in  order  that  it  may  produce  such  an  animal  as  man?" 


338  The   Unity  of  the  Organism 

the  subtler  aspects  of  the  problem.  The  little  we  shall  do 
in  this  way  may  be  introduced  by  the  query,  what  reason 
is  there  for  including  in  our  hypothesis  the  supposition  that 
it  is  "some  substance  in  the  air,  almost  certainly  oxygen," 
with  which  the  organism  reacts  chemically,  to  produce  con- 
sciousness and  all  other  phenomena  of  life?  Why  single  out 
this  substance  from  the  other  elementary  substances  essential 
to  life,  as  for  instance  carbon  or  nitrogen?  *  M}^  reply  be- 
gins by  recalling  the  immemorial  recognition  of  the  "breath 
of  life"  the  "life  giving  air"  and  so  on,  of  universal  experi- 
ence. It  is  well  to  recall  likewise  such  semi-philosophic  con- 
ceptions as  that  of  the  pneuma  or  "psychical  breath  of  life" 
of  later  Greco-Roman  philosophy.  The  inextricable  en- 
tanglement, historically,  of  breath  and  air  with  spirits  is 
also  worth  remembering,  especially  the  continuance  of  this 
into  the  modern  period  of  scientific  analysis,  unmistakeable 
traces  of  which  are  seen  in  the  writings  of  William  Harvey 
and  the  foremost  physiologists  of  the  era  to  which  he  be- 
longed. For  example,  the  spiritus  nitro-aereus  of  John 
Mayow  which,  we  now^  know,  was  his  term  for  oxygen  as 
glimpsed  first  in  the  history  of  science,  may  be  mentioned. 

More  important  than  any  of  these  reminders  from  the  his- 
tory of  knowledge  is  that  of  the  familiar  fact  that  the  most 
crucial  evidences  of  truly  independent  or  autonomous  life  of 
the  individual  higher  animal  are  respiratory.  That  the  new 
born  human  babe's  first  breathing-act  is  its  first  genuine  in- 
dependent life-act  is  one  of  the  most  commonplace  of  truths. 
And  recall  how  the  "return  of  life"  as  we  say  of  the  nearly 
drowned  person,  and  of  one  who  has  "fainted  dead  away" 
is  marked  by  the  resumption  of  respiratory  activities.  Cer- 
tain reflexes,  as  those  from  stimulating  the  eyelids,  and  pos- 

*  The  argument  in  answer  to  this  query  should  be  taken  as  an  exten- 
sion of,  and  in  important  respects  a  replacement  of,  that  contained  in 
my  essay.  Is  nature  infinife?^''  wherein  I  discuss  the  specificity  of  in- 
dividual' organisms  as  indicated  by  how  they  use  their  nutrient  sub- 
stances. 


Slefch   of  an  Off/a iiismal  Theory  of  Consciousu^ss      iV.V.) 

sibly  certain  heart  flutteriiigs,  may  Ik-  mhhv  iKTsisteiit  m.jvc- 
ments  than  those  connected  with   hrcathinn-.      Uui    these  are 
less  certain  signs  of  individual  JiiV-.     It  is  only  to  philosopliy 
of  the  elenientalist  sort  that  the  im-rc  twitch  of  a  hand  or  an 
eyelid  or  a   trace  of  heart   action   would    he   a   satisfactory 
proof  of  life.     Nor  would  it  be  to  a  philosoj)her  of  this  school 
should  the  "living  substance"  under  observation  haj)])en   to 
pertain  to  a  loved  relation  or  friend.   Satisfactory  evidence 
of  life  in  this  case  would  come  only  with   the  nearly  siinul- 
taneous    return    of    breathing   and    consciousness.      A    ri"-ht 
interesting   section    could    be    written    at    this    })oint   on    the 
importance  of  nutriment  in  the  ordinary  sense,  and  of  drink, 
as  compared  with  air  at  the  very  beginning  and  ending  stages 
of  the  individual  life.      For  instance   such   questions   would 
have  to  be  considered  as  that  of  the  independence  of  the  new 
individual   for   a  while  at   the  outset   on   food-yolk   in   many 
animals  below  the  mammals,  and  on  ])lacental  connections  in 
mammals ;   that   is   on  material  metabolically   elaborated    by 
the  older  or  parent  individual.     But  such   a  discussion  not 
being  indispensable  to  this  sketch,  must  be  foregone.    Enough 
here  to  emphasize  the  fact  that  while  it  may  be  entirely  jus- 
tifiable to  regard  oxygen  as  a  food  as  some  good  modern 
physiologists  do  the  two  important  facts  should  never  be  lost 
sight   of  that   (1)    oxygen   (air)    is   the  one  and   only   ever- 
present  and  never  varying  constituent   of  the  dietary.      In 
other  words  that  it  is  the  one  constituent  which  nature  sup- 
plies as  by  "free  grace"   to  use  a  good  old   theological  ex- 
pression; and  that  (2)  oxygen  is  the  one  and  only  food  that 
needs  no  digesting  and  so  no  digestive  organs  or  tissues  set 
apart  for  its  metabolic  elaboration.* 

Oxygen  is  the  only  food  which  ])asses  dlnctlv  as  such   to 

*  Were  the  view  held  hy  some  physiologists,  that  the  al\e«>lar  rjiithe- 
liiini  of  the  lungs  traiisinits  atiiiosplu-iic  oxygt'ii  to  tin-  liiood  l.y  an  i«ti\«* 
proeess  spoken  of  as  a  secreting,  this  stalcnu-nt  wonltl  nerd  iiiodifving 
somewhat.  However,  the  view  does  not  seem  t<»  he  aeeejjted  I'v  most 
authorities. 


340  The  Unity  of  the  Organism 

every  part  of  the  organism.  In  oxygen  the  organism  finds 
one  of  its  most  fundamental  food  materials  for  which  it  does 
not  normally  have  to  go  in  search  or  to  compete  with  other 
organisms.  The  familiar  fact  and  its  significance  appear  not 
to  have  attracted  the  attention  of  biologists  much.  Even 
L.  J.  Henderson  ^^  who  has  written  so  illuminatingly  on 
many  aspects  of  organic  adaptiveness  says  nothing  definite 
on  this  point.  These  two  facts  are  weighty  reasons  for  my 
proposal  to  look  upon  oxygen  as  one  chemically  elementary 
substance  and  the  organism  as  another,  the  reaction  be- 
tween which  is  basal  in  the  production  of  consciousness  and 
all  life  phenomena.  Consequently  the  problem  of  how,  ex- 
actly, the  organism  endowed  with  full-fledged  consciousness 
reacts  toward  oxygen  is  certainly  one  of  the  most  important 
of  all  problems  on  the  purely  physico-chemical  side  of  life. 
And,  as  said  early  in  this  sketch,  it  is  just  here  that  my  the- 
ory is  most  avowedly  hypothetical.  It  would  be  quite  out  of 
the  question  to  present  in  the  remaining  pages  of  this  book, 
even  had  I  the  requisite  knowledge  for  doing  so,  all  that 
might  profitably  be  said  on  the  subject.  Consequently  only 
two  or  three  of  what  seem  to  me  the  most  crucial  matters  will 

be  mentioned. 

In  the  first  place  I  ask  the  reader  to  recall  what  has  been 
said  in  various  of  the  preceding  chapters  which  have  brought 
out  the  indubitable  trend  of  the  interpretation  of  life  phe- 
nomena according  to  the  principles  of  physical  chemistry, 
away  from  the  elementahstic  conception  of  the  organism. 
The  interpretation  of  the  organic  cell  as  a  system  of  phases 
in  dynamic  equilibrium,  so  strongly  set  forth  by  Hopkins 
and  Bayless  will  be  remembered.  And  this  will  call  to  mind 
the  sharp  way  in  which  the  new  conception,  with  its  appeal 
to  the  role  of  surface-layers,  membranes,  and  areas  of  con- 
tact between  all  sorts  of  constituent  substances,  sets  itself 
over  against  such  pseudo-objective  conceptions  as  that  of 
biogens,  not  to  mention  the  horde  of  out  and  out  subjectivis- 


Sketch  of  an  Organismal  Theory  of  Consciousness      li-il 

tic  "elements"  of  which  paiigeiis  and  deterininuiils  Imve  per- 
haps liad  the  greatest  vogue.  The  importance  of  tlie  unti- 
eJenientahstic  tendency  of  physical  chemistry  when  it  comes 
to  be  applied  to  biological  ])roblems  is  greatly  enhanced,  it 
appears  to  me,  by  the  circumstance  that  J.  Willard  (Jibbs, 
who  was  one  of  the  very  first  to  appreciate  in  a  full  scientific 
sense  the  importance  of  massive  as  contrasted  with  minute- 
particle  phenomena  in  inorganic  nature,  and  so  was  one  of 
the  "fathers"  of  i)hysical-chemistry,  made  no  assumptions 
about  the  invisible  composition  of  substances  in  his  treat- 
ment of  "Heterogeneous  Equilibrium"  and  allied  topics. 
"Certainly,"  writes  Gibbs,  "one  is  building  on  an  insecure 
foundation  who  rests  his  work  on  hypotheses  concerning  the 
constitution  of  matter."  ^^  If  this  is  true  as  touching  the 
relatively  simple  structures  and  movements  in  the  lifeless 
world  how  much  more  obviously  true  is  it  as  touching  the  liv- 
ing world,  and  especially  such  life  phenomena  as  human  con- 
sciousness ! 

So  we  are  able  to  requisition  one  of  the  admittedly  most 
important  advances  of  modern  times  in  inorganic  science  as 
support  for  the  supposition  that  the  air  we  breathe,  and 
presumably  its  oxygen,  contributes  in  some  direct  and  funda- 
mental way  to  the  production  of  consciousness  even  though 
this  substance,  if  its  "ultimate  nature"  is  what  inorganic 
chemistry  and  physics  have  hitherto  attributed  to  it,  has  lit- 
tle or  nothing  to  suggest  that  it  possesses  such  a  unique 
latent  attribute.  The  reader  should  not  fail  to  recall  here 
Hume's  recognition  of  the  "secret  powers"  of  substances. 

But  is  it  not  possible  that  physico-chemical  and  physi- 
olop-ical  knowledge  of  oxvcren  and  air,  the  'M)reath  of  lifr," 
do  contain  somewhat  more  to  justify  tiie  suj)positi()n  than  is 
usually  recognized?  In  this  connection  I  relate  that  om-  of 
the  most  mentally  adhesive  statements  I  ever  heard  from  a 
blo-chemist,  its  adhesiveness  de]K>nding  largely  on  the  fait 
that  the  chemist  was  one  of  great  experience  as  a  laboratory 


^42  The  Unity  of  the  Organism 

investigator,  was  to  the  effect  that  chemical  analyses  make 
known  what  they  find  and  absolutely  no  more.  In  other 
words  such  analyses  never  exclude  the  possibihty  of  sub- 
stances other  than  those  found.  And  this  chemist  asserted 
furthermore  that  all  organic  analyses  leave  residues  to  some 
extent.  No  manipulative  methods  are  known,  it  appears, 
capable  of  effecting  a  really  complete  analysis  of  any  or- 
ganic substance.  Whether  these  restrictions  on  analyses 
still  hold  I  am  not  sure,  though  I  have  seen  or  heard  nothing 
which  leads  me  to  suppose  they  do  not. 

It  is  this  general  shadow  of  manipulative  imperfection 
which  overhangs  all  formal  physics  and  chemistry,  coupled 
with  the  advances  being  made  from  time  to  time  in  our 
knowledge  of  oxygen  and  air  which  has  led  me  to  put  into  my 
hypothesis  a  shade  of  doubt  as  to  whether  oxygen  is  the  con- 
stituent of  the  air  the  reaction  of  which  with  the  organism 
produces  consciousness.  The  demonstration  of  helium  and 
argon,  and  probably  neon,  crypton,  and  xenon  in  atmos- 
pheric air,  all  within  a  little  more  than  two  decades,  has 
influenced  my  thinking  in  the  same  direction.  Besides,  the 
idea,  become  a  commonplace  of  physics  and  chemistry  in  a 
single  night,  figuratively  speaking,  that  the  "atom  is  as  com- 
plex as  the  solar  system"  has  had  its  part  in  shaping  my 
conceptions ;  as  have  also  such  well-credentialed  conceptions 
from  the  inorganic  sciences  as  that  "Uranium  II"  is  "a  long- 
lived  element"  which  is  the  "parent  of  the  actinium  series  of 
elements,  but  has  no  genetic  connection  with  the  uranium 
series" ;  and  that  "in  the  lead  pleiad  there  are  seven  ele- 
ments having  quite  different  atomic  weights."  ^^ 

The  extent  to  which,  as  exemplified  by  this  case,  the  inor- 
ganic sciences  have  found  themselves  driven  into  the  organic 
realm  for  terms  with  which  to  express  their  new  conceptions 
must  impress  every  thoughtful  person.  Earlier,  what  we 
might  describe  as  p\4rely  contemporaneous  physical  dynamics 
had  to  borrow  such  terms   as  energy,  power,   force,  work. 


Sketch  of  an  Organismal  'riwor/i  of  Consrioustwss      343 

from  the  nomenclature  of  liviiio-  })cin^rs.  Iwiter,  with  tlie  per- 
meation of  all  knowledge  by  the  eoncej)tioM  of  the  natural 
or  derivative  origin  of  everything  (a  genuinely  organic  con- 
ception, notice),  has  come  even  for  elementary  cliemical  sub- 
stances, the  induction  into  i)hysics  and  chemist rv  of  such 
ideas  as  genetic  relations,  parenthood,  and  li-ngth  of  lifr. 
So  my  suggestion  tliat  the  air  we  breathe  nuist  be  rcco<rni/c(l 
to  possess  latent  attributes  which  by  reacting  with  the  or- 
ganism produce  consciousness,  falls  into  a  genetic  scries  in 
the  history  of  the  interpretation  of  nature. 

The  very  important  question,  as  already  indicated,  of  ex- 
actly how  atmospheric  or  molecular  oxygen  oj)erates  in  the 
living  being  generally  and  the  conscious  being  ])articularly, 
is  largely  for  the  future  to  answer.  One  should  never  fail, 
however,  to  couple  this  question  with  the  same  question  as  to 
the  behavior  of  oxygen,  and  for  that  matter  of  any  other 
chemical  substance,  in  any  reaction  whatever.  Exactly  how, 
for  example,  does  oxygen  operate  with  hydrogen  to  produce 
the  attribute  of  ref  rangibility  of  water ;  or  with  phosphorus 
to  produce  the  peculiar  glow  which  that  substance  may  ex- 
hibit under  some  conditions.'^ 

Concerning  the  positive  knowledge  and  the  views  as  to 
details  of  the  action  of  oxygen  in  connection  with  the  or- 
ganism, only  a  little  can  be  said  here  though  that  little  may 
be  very  important.  Looked  at  from  the  standpoint  of  the 
old,  the  orthodoxly  atomistic  chemistry,  probably  the  most 
anomalous  thing  about  my  hypothesis  is  that  the  organism 
conceived  as  equivalent,  chemically  s])eaking,  to  an  elenun- 
tary  substance,  is  the  unciuestioned  fact  that  the  organism  is 
not  only  composed  of  several  chemical  substancis,  but  that 
one  of  these  is  oxygen  itself.  Stated  iialdly,  the  anomaly  is 
that  two  chemical  substances  are  supposed  to  react  upon  each 
other,  one  of  which  (the  organism)  is  known  not  only  not 
to  be  simple,  but  to  contain  the  otlu  r  substance.  Hut  even 
the   old   chemistry   with    its   "compound    radicals,"    ot*   which 


344  The  Unity  of  the  Organism 

cj^anogen  (CN)2  is  said  to  have  been  the  first  discovered,  and 
of  which  the  unitedly-acting  combinations  of  carbon  and 
hydrogen  as  methyl,  CHg,  affords  some  slight  support  for 
our  conception  so  far  as  the  mere  matter  of  chemically  uni- 
tary compoundedness  is  concerned.  In  so  far,  however,  as 
technical  chemistry  can  be  drawn  upon  for  supporting  our 
hypothesis,  it  is  the  new,  or  physical  chemistry,  as  has 
been  repeatedly  stated,  that  is  our  main  reliance.  Unless  I 
am  greatly  deceived,  the  real  inwardness  of  that  great  move- 
ment in  inorganic  science  is  against  the  age-old  conception  of 
the  ultimate  adequacy  of  atoms  to  explain  inorganic  na- 
ture, almost  as  positively  as  the  organismal  conception  is 
against  the  ultimate  adequacy  of  any  constituent  element 
whatever,  to  explain  organic  nature.  The  surface  energies, 
for  example,  developed  at  contact  faces  and  giving  rise  to 
the  phenomena  of  adsorption  *  appear  to  be  not  a  whit  less 
real  and  ultimate  energies  than  are  any  that  can  be  attrib- 
uted to  atoms  and  molecules  taken  as  such.  And,  be  it  no- 
ticed, one  of  the  most  distinctive  things  about  these  areal  and 
massive  energies  is  that  they  dominate  atomic  and  molecular 
energies  to  a  certain  extent.  This  is  just  what  the  now  uni- 
versally recognized  principle  of  "mass  action"  is  in  so  far  as 
such  action  has  been  studied  enough  to  make  possible  its  for- 
mulation into  law ;  that  is  enough  to  learn  how  it  influences 
velocity  and  quantity  of  chemical  change.  But  would  any 
careful  physicist  or  chemist  pretend  to  know  to  a  certainty 
that  such  action  is  restricted  to  influence  of  that  sort.?  Surely 
not.     Are  we  certain  for  instance  that  it  can  not  under  any 

*  Adsoriition  is  the  loading  of  the  surface  of  a  solid  body  immersed  in 
a  solution,  with  the  dissolved  substance.  Thus  it  is  by  adsorption  that 
charcoal  takes  the  coloring  matter  out  of  a  colored  solution.  The  action 
results  from  the  facts  that  there  is  surface  tension  at  the  interfaces  be- 
tween the  charcoal  and  the  liquid,  and  that  this  tension  is  lessened  by 
the  presence  of  the  dissolved  color-substance  in  the  liquid.  The  sub- 
stance then  moves  to  the  place  of  lessened  tension  and  concentrates  on 
the  surface  of  the  solid.^-  The  principle  has  very  wide  application  in 
nature,  particularly  in  organic  nature,  where  colloidal  substances  and 
water  are  in  contact  so  extensively. 


Sketch  of  an  Orgamsmal  Theory  of  ConsciousTwss     S45 

circumstances  influence  qualitative  as  well  as  quantitative 
change?  Surely  we  are  not.  This  of  course  is  far  from 
contending  that  mass  action  actually  does  influence  quali- 
ties. My  sole  point  is  that  so  long  as  there  is  lack  of 
certainty  that  it  does  not  or  may  not  exert  such  influence 
any  assumption  which  implies  such  certainty  is  unwarranted 
and  unscientific. 

Putting  together,  then,  the  physically  massive  conce])- 
tlons  of  inorganic  chemistry  and  the  organismal  conceptions 
of  bio-chemistry  what  seems  to  follow  touching  the  chemico- 
substantive  composition  of  organisms  is  that  a  portion  of 
all  the  substances  essential  to  life,  carbon,  oxygen  and  others, 
have  been  combined  from  all  eternity  (whatever  Ije  the  mean- 
ing of  the  phrase)  in  the  peculiar  way  called  organic,  while 
other  portions  have  remained  in  the  state  called  inorganic. 
This  leads  me  to  remark,  quite  incidentally  so  far  as  this 
discussion  is  concerned,  that  according  to  this  view  the  as- 
sumption would  be  that  organisms  have  always  existed,  or  at 
least  that  they  have  existed  as  long  as  "matter"  or  anything 
else  of  which  we  have  any  information  or  clear  conception, 
has  existed.  The  warrantableness  of  this  assumption  I  am  re- 
lieved from  arguing  here  from  having  treated  the  problem 
at  some  length  in  another  place.  (Ai'e  xce  obliged  to  suppose 
the  spontaneous  generation  of  life  ever  occurred?Y^  All 
that  need  be  said  now  about  the  outcome  of  that  discussion  is 
that  the  warrantableness  lies  in  the  absence  of  any  ground 
for  assuming  the  contrary.  I  take  my  position  squarely  on 
the  direct  evidence  in  the  case.  All  the  evidence  of  that  sort 
we  have — and  in  that  discussion  I  emphasize  the  fact  of  its 
vast  quantity — is  to  the  effect  that  organisms  are  produced 
by  other  organisms  known  as  parents  and  in  no  other  way.* 

*  To  the  stock  and  rather  vapid  rejoinder  that  sneli  a  sohitioii  of  tlie 
j)rol)h'ni  of  tlie  orifrin  of  life  is  no  solution  at  all,  i»nt  only  a  jMittitifr  t)lTof 
the  dittieulty,  the  ohvious  reply  from  my  standpoint  is  that  I  am  mnkinj; 
no  pretense  of  "solvinji;  the  prohlem,"  as  "solution"  would  he  meant  in 
the  anticipated  rejoinder.  From  my  standpoint,  however,  the  everlast- 
ingly-from-parents  hypothesis  woidd  he  a  solution  of  tin-  prohlem  if  the 
hypothesis  were  proved  true. 


^46  The  Unity  of  the  Organism 

We  can  now  state  briefly  as  much  more  of  the  bio-chemical 
aspect  of  the  problem  as  seems  indispensable  to  our  present 
argument.    A  few  remarks  on  what  the  physiology  of  our  day 
often  calls  tissue  respiration  will  compass  what  is  in  mind. 
The  key  fact  in  this  is  of  two-fold  character:  (1)  The  tissues 
of  the  organism,  not  its  blood  or  any  other  fluids,  contain  the 
substance  which  is  in  the  strictest  sense  living.      (2)   This 
substance  is  called  living  because  chemical  changes  of  a  very 
distinctive  sort  are  going  on  in  it.     These  changes  are  of  a 
fundamentally    double    nature    as    regards    atmospheric    or 
molecular   oxygen;   namely,   combinative   and  incorporative 
change,  and  separative  and  expulsive  change.     The  last-men- 
tioned, the  separative  and  expulsive  change,  is  known  as  oxi- 
dation and  manifests  itself  to  ordinary  experience  in  the  dis- 
charge of  oxygen  combined  with  carbon  as  carbon  dioxide, 
and  in  the  setting  free  of  energy  in  the  form  of  muscular  and 
other  work,  and  of  heat.     The  first-mentioned,  or  incorpora- 
tive change,  consists  in  taking  in  and  storing  up   oxygen, 
"somehow,"  as  the  more  carefully  worded  physiologies  put 
it.     This  statement  may  be  taken  as   a  very  brief  natural 
history  description  of  the  most  fundamental  steps  in  what 
formal  physiology  calls  metabolism  with  its  two  aspects,  the 
constructive,  or  anabolic,  and  the  destructive,  or  katabolic. 
Probably  no  one  will  question  that  this  conception  of  the 
foundations  of  the  life  process  for  nearly,  if  not  quite,  all 
animal  life  is  that  held  by  the  best  physiologists  since  the 
time  of  C.  Bernard  at  least.     No  physiologist  whom  I  have 
consulted  has  stated  the  nature  of  the  process  more  definitely 
than  has  Sir  Michael  Foster.     "The  Respiration,"  he  writes, 
"of  the  muscle  then  does  not  consist  in  throwing  into  the 
blood  oxidizable  substances,  there  to  be  oxidized  into  car- 
bonic acid  and  other  matters ;  but  it  does  consist  in  the  as- 
sumption and  storing  up  of  oxygen  somehow  or  other  in  its 
substance,  in   the  building  up  by  help   of  that  oxygen   of 
explosive  decomposable  substances,  and  in  the  carrying  out 


Sketch  of  an  Orc/atiismal  Thcorij  of  Consciousness      847 

of  decompositions  whereby  carbonic  acid  and  other  matters 
are  discliarged  first  Into  the  siil)stance  of  the  nuiscle  and 
subsequently  into  the  bk)od."'''  And  lie  points  out  in  other 
connections  that  what  is  true  of  nuiscle  in  this  reL^•l^d  is  es- 
sentially  true  of  all  other  tissue  systems.  In  another  still 
more  recent  text  book  we  read:  "Nothing  definite  is  kiiown, 
however,  as  to  the  nature  of  the  ]u*obable  combinaticms 
formed  by  oxygen  with  the  different  materials  for  l)uilding 
up  muscles  and  other  tissues,  or  of  the  intermediate  anabolic 
and  katabolic  forms  througli  which  it  passes  in  combining 
with  carbon  into  carbonic  acid."  '^^  And  this  autlior  then 
expresses  what  arc,  apparently,  his  own  views,  by  quoting 
from  Foster  as  follows:  "The  whole  mvsterv  of  life  lies  liid- 
den  in  the  story  of  that  progress  [that  of  construction  and 
destruction  in  the  tissues]  and  for  the  present  we  nnist  Ik? 
content  with  simply  knowing  the  beginning  and  the  end." 

The  kernel  of  my  suggestion  so  far  as  metabolism  is  con- 
cerned, is  that  the  anabolic,  or  the  assimilative,  the  trulv 
synthetic  aspect  of  the  com})lete  operation,  is  the  contimial 
renewal,  or  keeping  up  of  the  oxygen  constituent  of  the 
organism  which  comes  to  it  by  heredity,  that  is  which  has 
always  been  in  the  "line  of  descent."  It  is  the  maintenance 
of  what  might  be  spoken  of  as  the  original  oxygen  constitu- 
ent of  the  organism.  There  would  always  then  be  operating 
in  the  organism  oxygen  of  two  sources,  that  from  tlu"  one 
source  designated,  employing  our  well-established  evohitional 
terminology,  phylogenic  or  hereditary  oxygi'n  :  and  the  other 
ontogenic  or  individual  oxygen.  In  general  the  same  kind 
of  reasoning  would  hold  for  the  other  chemical  simples,  car- 
bon, nitrogen,  and  so  on;  but  these  are  in  (julte  a  difrirint 
status  from  oxygen  owing  to  thi-  fact  that  they  are  not 
normally  taken  by  the  animal  organism  in  tlie  pure  or  uncom- 
bined  state,  but  only  in  some  other  organic  combination, 
as  food  in  the  ordinary  sense. 

Metabolically  ex]iressed,  then,  we  may  say  in  short   that 


348  The  Unity  of  the  Organism 

the  warrantableness  for  considering  the  individual  organism 
as  a  chemical  element,  is  the  fact  that  it  maintains  its  identity 
as  regards  all  its  elementary  constituents  except  one,  oxygen, 
be  wrenching  these,  so  to  speak,  from  other  organic  com- 
pounds (by  digesting  these)  and  then  by  synthesizing  the 
elements  into  its  own  particular  substance.  Another  way  of 
expressing  the  same  conception  is  to  say  that  the  organism 
is  an  element,  chemically  speaking,  because  it  reacts  directly 
in  a  chemical  sense  with  another  element. 

Did  this  chapter  pretend  to  be  anything  more  than  a 
sketch  of  a  theory  of  consciousness  a  considerable  discus- 
sion of  the  "activation"  of  oxygen  would  naturally  come  in 
somewhere,  perhaps  at  this  point.  The.  essense  of  activation 
is  the  fact  that  when  oxygen  passes  into  the  organism  by  the 
respirator}^  process  it  is  somehow  changed  into  a  condition 
which  enables  it  to  oxidize  living  tissue-substances  as  it 
can  not  to  any  degree,  seemingly,  when  brought  into  con- 
tact with  the  same  substances  outside  the  organism.  This 
discussion  would  involve  the  various  theories  which  have 
been  put  forward  to  account  for  this  phenomenon,  as  those 
which  make  use  of  the  principle  of  enzymes,  of  peroxides  or 
of  some  other.  All  that  our  aims  here  require  us  to  notice 
is  that  nothing  conclusive  as  touching  the  nature  of  activa- 
tion would  come  from  the  discussion.  How  unsatisfactory 
a  state  this  whole  subject  is  in  may  be  seen  from  the  follow- 
ing words  of  a  foremost  American  biochemist:  "It  has  been 
a  popular  practice  to  appeal  to  hypothetical  enzymes  to 
explain  some  of  the  obscure  chemical  transformations  in  the 
organism.  Thus  we  have  been  wandering  through  the  mazes 
of  the  oxidases,  oxygenases,  peroxidases,  reductases,  cata- 
lases  and  other  products  of  perplexing  nomenclature  in  the 
hope  of  escaping  the  uncertainties  of  intermediary  meta- 
bohsm."  3« 


Sketch  of  an  Organismal  Theory  of  Cansciou^mss      3i9 

Summcd-up   Statement   of  Justification   of    tlu-   Jl/jpothcun 

The  final  gathcrliifr-up-.-ind-puttincr-to^athcr  may  now  be 
made  of  all  that  has  lx?en  said  about  the  physico-chemical 
aspect  of  the  organism  on  the  one  hand,  and  alx>ut  its  j)sy- 
chical  aspect  on  the  other.  That  is  to  say,  we  are  now  roady 
to  epitomize  the  results  of  our  examination  of  the  ancient 
and  honorable  but  withal  unsolved  jjroblcm  of  liow  Body  and 
Soul  go  together.  As  regards  "body"  or  "the  physical"  we 
have  been  led  to  the  physico-chemical  conception  of  the  or- 
ganism as  a  well-nigh  inconceivably  complex  mass  of  sub- 
stances, mostly  in  the  colloidal  state,  operating  as  a  system 
of  phases  in  dynamic  or  constantly  changing  equilibrium. 
As  regards  "soul"  or  "the  psychical,"  we  have  found  also  a 
series  of  phases  of  activities,  namely  tlie  phases  of  intellect 
and  reason,  those  of  instinct,  those  of  feeling  and  emotion, 
those  of  the  will,  those  of  the  tropisms  and  the  "simple  re- 
flexes," and  finally  those  of  simple  protoplasmic  res])onse. 
According  to  my  hypothesis,  the  phases  of  the  })io-chemico- 
physical  sort  and  the  phases  of  the  psychical  sort  have  com- 
mon ground  in  the  organism  as  a  whole,  the  phases  of  in- 
tellect and  reason  corresponding  to  the  cerebro-spinal  nerv- 
ous system;  the  phase  of  instinct  corresponding  probably  to 
the  autonomic  nervous  system ;  the  phases  of  feeling  and  cnio- 
^  tion  corresponding  mainly  to  the  glandular  and  visceral  sys- 
tems;  those  of  the  will  to  the  body-muscular  system;  those 
of  the  tropisms  and  simple  reflexes  to  the  receptor-conduct or- 
efFector  systems;  and  finally  those  of  simj)le  protoplasmic 
response  to  the  fundamental  protoplasmic  mechanism  of 
response,  whatever  its  structure. 

According  to  the  scheme  presented  in  the  sketcli  ami 
summed  up  here,  just  as  physical  functioning  and  physical 
form  reach  back  to  the  very  dawning  of  animal  life,  both  in 
the  individual  and  in  the  race  or  type,  so  consciousness  with 
its  nether  limits  in  what,   following  the  ternn'nology  of  em- 


350  The  Unity  of  the  Organism 

bryology  (see  section  on  the  pro-morphology  of  the  egg- 
cell,  Chap.  8),  might  be  called  pro-consciousness,  is  an 
attribute  of  all  animal  organisms.  As  comparative  anatomy 
and  physiology  have  made  us  familiar  with  the  physical  as- 
pect of  the  animal  organism  existing  as  the  fully  realized 
or  developed  adult  at  one  end  of  the  ontogenic  series,  and  as 
the  unrealized  adult  or  germ  at  the  other  end  of  the  same 
series,  exactly  so  its  psychology  gradually  familiarizing  us 
with  the  realized,  or  adult  mind  at  one  end  of  the  ontogenic 
series,  and  as  the  unrealized  or  germinal  mind  at  the  other 
end  of  the  same  series.  When  we  affirm  that  the  completed 
individual  organism  is  latent  in  the  germ,  we  must  under- 
stand that  the  psychical  aspect  no  less  than  the  physical 
aspect  is  so  latent.  With  ver}^  little  doubt,  it  seems  to  me, 
the  real  meaning  of  the  so-called  sub-conscious,  and  of 
psycho-analysis  as  a  method  of  investigating  it,  is  that  the 
ontogenic  stages  of  the  psychic  life  of  the  human  organism 
are  being  discovered  and  that  a  method  of  investigating  these 
stages  is  being  worked  out.  Freud  and  his  followers  have 
been  and  still  are  somewhat  in  the  dark,  I  think,  as  to  just 
what  they  are  doing,  albeit  their  discoveries  and  methods  are 
of  the  utmost  importance. 

REFERENCE  INDEX 
1.  Wheeler,  W.  M 519       21.  Ritter,  W.  E.  (1918==)   Essay  2 

3.  Sedewick  &  Taylor 262-3       22.  Huxley,   T.   H 98 

4.  Rovce,  J 2       23.  Huxley,   T.   H 96 

5.  Royce,  J 3       24.  Huxley,  T.  H 100 

6.  James,  Wm 1-341       25.  Huxley,  T.  H 94 

7.  Wheeler,  W.  M 521       26.  Dewey,  John    9 

8.  Whitman,  CO 310       27.  Darwin,  Chas 

9.  James,  Wm n-451       28.  Crile,  G.  W 

10.  Cannon,   W.    B 280       29.  Morgan,  C.  Lloyd   256 

11.  Cannon,  W.  B 282       30.  Henderson,  L.  J 

12.  James,  Wm 11-450       31.  Gibbs,  J.  W Pref.  x 

13.  James,  Wm n-451  32.  Lewis,   Wm.   C.   McC.  .  .H,  303 

14.  Hopkins,   F.   G 217  33.  Ritter,  Wm.  E.  (19180  Essay  1 

15.  Hopkins,  F.  G 220      34.  Foster,  M 469 

16.  Hopkins,  F.  G 220       35.  Luciani,  L I,  395 

17.  Sellars,  R.  W 75       36.  Mendel,  L.  B 21 

18.  Descartes,  Rene..Pt.  1,  Sec.  8  37.  Ritter,    Wm.    E.    (1918==) 

19.  Montague,  W.  P 120       38.  Wendt,  G.  L 442 

20.  Hume,  David. .  .Sec.  IV,  Pt.  1       39.  Stout,   G.    F 680 


POSTSCRIPT 

THE  argument  in  favor  of  the  organismal  way  of  viewing 
living  nature  has  now  run  what  appears  to  mc  its  natural 
course,  to  its  inevitable  end.  Yet  I  cannot  ])ring  myself  to 
write  "Finis"  without  making  a  few  remarks  which  thougli 
connected  vitally  with  the  argument,  do  not  seem  an  essential 
part  of  it. 

These  remarks  concern  the  general  effect  of  the  organismal 
standpoint  on  those  who  may  grasp  it  firmly  and  adopt  it 
unreservedly.  Since,  as  pointed  out  in  the  "Historic  Back- 
ground" with  which  this  book  opens,  the  standpoint  has  been 
recognized  by  biologists  with  varying  degrees  of  fullness 
from  the  time  of  Aristotle  at  least,  there  can  be  no  doubt 
that  the  human  mind  is  naturally  attuned,  as  one  might  say, 
to  this  general  type  of  response  to  organic  phenomena.  It 
seems  therefore  fitting  that  a  presentation  like  that  which  I 
have  made  should  be  accompanied  by  a  few  words  on  the 
probable  influence  of  a  wide  prevalence  of  the  organismal 
view.  The  pertinent  question  will  be  asked,  how  could  it 
have  come  to  pass  that  if  the  standpoint  has  been  so  long 
in  the  w^orld  it  should  have  missed  full  recognition  and  have 
failed  to  exert  its  due  influence?  The  reply  is  obvious  to  an 
attentive  reader  of  this  book:  At  no  time  until  the  present 
in  the  long  historical  growth  of  knowledge  of  the  living  world 
has  information  been  sufficient  to  make  possible  a  rounded- 
out  statement  of  the  conception.  To  illustrate,  it  is  only  in 
the  very  last  years  that  enough  has  been  known  of  the 
physical  chemistry  of  the  cell  to  engender  such  an  interpre- 
tation of  this  exceedingly  im])ortant  biological  entity  as  that 
which  biochemists  are  just  now  reaching.     Yet  this  interpre- 

351 


352  Postscript 

tation  is  indispensable  to  anything  even  approaching  a  full 
development  of  the  organismal  view. 

But  nothing  stands  out  more  boldly  from  the  pages  of 
this  book  than  the  insufficiency  even  yet,  of  actual  knowledge 
for  making  the  standpoint  complete.  If  therefore,  I  append 
to  my  presentation  a  brief  reference  to  the  larger  effect  the 
view  has  had  on  myself,  and  on  this  basis  forecast  what  the 
effect  would  be  on  thinking  people  generally  were  they  to 
make  it  their  own,  such  a  forecast  will  surely  be  in  harmony 
with  the  larger  purpose  of  the  book,  even  though  the  antici- 
patory remarks  have  no  place  in  the  presentation  itself. 

The  long  and  laborious  gathering  and  arranging  of  facts, 
and  weighing  of  natural  evidence  and  formal  arguments 
which  has  constituted  the  development  of  the  standpoint  in 
my  own  mind,  has  compelled  me- to  re-examine  and  re-assess 
the  whole  frame  and  fabric  of  my  spiritual  life.  Nothing,  so 
far  as  I  can  tell,  has  escaped.  Not  my  scientific  knowledge 
alone — my  professional  stock-in-trade — but  all  my  ideas  and 
beliefs  touching  religion,  art,  society,  politics,  industry,  per- 
sonal relations,  and  private  living,  have  come  in  for  their 
share  of  scrutiny  and  renovation. 

An  exceedingly  brief  "synoptic"  classification  and  char- 
acterization *  of  the  entire  range  of  these  effects  can  be 
given  in  the  terms  of  formal  science  and  philosophy. 

As  to  classification,  the  effects  fall  into  a  two-fold  group- 
ing. One  of  the  groups  appertains  to  the  great  province  of 
the  nature  of  knowledge;  the  other  to  the  equally  great 
province  of  the  nature  of  morals. 

The  characterization  of  effects  on  the  nature  of  knowl- 
edge which  seems  to  me  most  inclusive  and  most  practically 
significant,  may  be  stated  thus :  By  the  validation  of  ob- 
jective knowledge,  largely  through  the  principle  of  what  I 
have  called  standardization  of  reality,  but  partly  through 

*  See  my  essay,  The  Place  of  Description,  Definition  and  Classifica- 
tion (Ritter,  1918). 


Postscr'qyt  15.53 

the  organisnial  livpotht'sls  of  consciousness,  sucli  knoulcd^c 
is  elevated  to  tlie  rank  of  strict  e(|ualitv  with  "pure  tliou^ht," 
often  so-called;  that  is,  witli  suhjcctivc,  oi-  intiiltivi-  knowl- 
edge. In  this  way  niatheniatico-niechanistic  science  is  de- 
prived of  the  regal  place  it  has  claimed  for  itself  since  the 
era  of  Descartes  and  Leibnitz,  and  is  brought  to  the  plane 
of  absolute  equality  as  to  importance  and  dignity,  with 
sense-experiential  science,  liy  thus  adjusting  the  claims  of 
these  two  great  realisms  of  science,  an  attitude  toward  t  he 
infinite  totality  of  nature,  and  a  methodology  for  interpret- 
ing it,  which  have  hitherto  borne  the  stanij)  of  subjection 
and  inferiority  assume  their  rightful  places  in  the  great 
hierarchy  of  philosophical  science.  This  leveling-down  of 
mathematical  mechanics  and  the  deductive  method  and  level- 
ing-up  of  observational  knowledge  and  the  inductive  metljod, 
implies  the  complete  overthrow  of  psycho-physical  dualism 
in  psychology,  and  the  rescue  of  })ersonality  from  bondagi 
to  a  theoreticallv  infinite  monotony  of  "Matter  and  Knertrv." 

The  characterization  of  the  effects  of  the  organismal  \  i(  w 
on  morals  centers  around  the  perce])tion  that  in  the  establish- 
ment of  human  personality  the  persons  are  organically  in- 
terdependent upon  one  another;  that  is,  interde|)en(ii'nt 
through  their  "attributes  of  relation,"  this  resulting  in  the 
incorporation  of  men  into  a  ])luralistic  universe  far  more 
real  and  vital  than  })hiloso])hic  })luralism  has  hitherto  hreii 
in  position  to  grasj).  I'hrough  a  type  of  human  conduct 
guided  by  knowledge  of  these  principles  of  personality  and 
the  interdependence  of  personalities,  and  thiough  su)»|)le- 
menting  mathematico-mcchanistic  methods  of  study  by  a 
rigid  application  of  observational  and  statistical  methods,  a 
genuine  science  of  morals,  both  theoretical  and  practical, 
is  made  attainable. 

That  my  enterprise  of  developing  the  organismal  \  iiw  is 
only  part  and  ])arcel  of  the  gem  ral  current  of  iiitt  rj)retat ion 
of  living  nature  which  has  flowed  through  the  centuries  seems 


354  Postscript 

clear  even  from  my  meager  acquaintance  with  the  history  of 
philosophic  thought.  Thus  we  read  in  Windelband  {A  His- 
tory of  Philosophy,  Eng.  by  Tufts,)  :  "For  the  decisive  fac- 
tor in  the  philosophical  movement  of  the  nineteenth  century 
is  doubtless  the  question  as  to  the  degree  of  importance 
which  the  natural-science  conception  of  phenomena  may 
'  claim  for  our  view  of  the  world  and  life  as  a  whole."  (624). 
Then  after  speaking  of  the  sharp  antithesis  between  the 
W eltmischauung  elaborated  by  the  "Highly  strained  idealism 
of  the  German  Philosophy"  of  the  early  nineteenth  century, 
and  the  "materialistic  Weltanschauung''  of  the  later  decades 
of  the  same  century,  the  author  writes :  "If  we  are  to  bring 
out  from  the  philosophical  literature  of  this  century  and 
emphasize  those  movements  in  which  the  above  characteristic 
antithesis  has  found  its  most  important  manifestation,  we 
have  to  do  primarily  with  the  question,  in  what  sense  the 
psychical  life  can  be  subjected  to  the  natural-science  mode 
of  cognition."     (p.  6^5). 

That  Part  II  of  this  book  of  mine,  especially  Chaps.  20 
to  24,  go  a  long  way  toward  answering  the  cardinal  question 
formulated  by  Windelband  appears  to  me  certain.  And,  I 
may  add,  it  also  seems  quite  clear  to  me  that  the  gigantic 
struggle  at  arms  which  that  philosopher's  nation  has  now 
brought  upon  the  world,  is  one  of  the  strongest  proofs  that 
philosophic  thought  and,  following  this,  social  and  political 
leadership  in  Germany  have  failed  miserably  to  discover  the 
Via  Media  between  the  Weltanschauung  of  the  "highly 
strained  idealism  of  the  German  Philosophy"  and  the  mate- 
rialistic Weltanschauung  which  has  finally  reached  its  nat- 
ural climax  in  militaristic  brutism,  and  is  almost  certainly 
(Sept.,  1918)  approaching  its  overthrow. 

Nothing  could  more  fittingly  end  this  book,  devoted  as  it 
is  to  demonstrating  the  operative  nature  of  organic  unity 
in  one  of  its  great  segments,  than  a  reference  to  the  fact 
that  the  philosophy  of  life  now  determining  German  morals, 


Postscript  355 

and  which  has  drawn  its  in.s{)irali()ii  hirgcly  fiuni  the  liypo- 
thesis  of  natural  selection,  has  failed — patlictically  Ix'vond 
the  power  of  words  to  express  if  done  unintentionally;  and 
criminally  in  equal  measure  if  done  intentionally — to  under- 
stand the  real  meaning  of  Darwin's  teaching  as  a  whole. 

Certain  it  is  that  had  the  German  philoso])hers  of  Mdclit- 
politik  recognized  the  place  of  uiujualifii'd  suj)remacy  as- 
cribed by  Darwin  to  the  mental  and  moral  endowments  of 
man,  it  would  have  been  impossible  for  them  to  make  the 
dogma  of  survival  of  the  fittest  serve  their  ends  in  any  such 
way  as  they  have  made  it,  and  done  so  honestly.  Attentive 
reading  of  The  Descent  of  Man  makes  it  perfectly  plain 
that  Darwin  simply  accepted  all  the  higher  human  attri- 
butes— moral,  esthetic,  and  religious,  no  less  than  those  of 
the  intellect — as  fundamental  data  in  his  reasoning  about 
man's  evolution.  His  sole  effort  as  touching  these  was  merely 
to  see  in  how  far  they  could  be  regarded  eitlu  r  as  helped 
forward  in  their  development  by  natural  selection,  or  at  least 
as  not  inconsistent  with  it.  Apparently  it  never  even  oc- 
curred to  him  to  regard  his  hypothesis  as  supreme-over-all, 
so  that  all  attributes  whatever,  the  noblest  ones  of  man  with 
the  rest,  must  either  be  forced  into  conformity  with  it,  or 
their  reality  and  power  virtually  denied.  "I  fully  subscribe 
to  the  judgment,"  runs  the  opening  sentence  of  the  chapter 
on  "The  Moral  Sense,"  etc.,  "of  those  writers  who  maintain 
that,  of  all  the  differences  between  man  and  the  lower  ani- 
mals, the  moral  sense  or  conscience  is  by  far  the  most  im- 
portant." And,  especially  significant  at  this  time,  Darwin 
quotes  with  obvious  a])j)r()val,  an  apostrophe  to  Diit\  hy 
Kant,  in  which  this  "Wondrous  thought"  is  represented  as 
"holding  up  its  naked  law"  in  the  soul,  and  demanding 
reverence.  Darwin's  entire  discussion  in  this  ])art  of  tjie 
Descent  makes  it  clear  that  what  he  had  in  mind  was  to 
discover  as  far  as  possible  the  germs  of  "conscience,"  of 
"feeling  of  right  and  wrong,"  of  an  "inward  monitor,"  of 


356  Postscript 

"sympathy,"  of  "parental  and  filial  affection,"  of  "social 
affection,"  of  the  "instinct  of  self-sacrifice"  and  so  on,  in  the 
lower  animals  so  as  to  have  a  starting  point  for  these  attri- 
butes as  they  occur  in  civilized  man.  It  was  not  at  all  his 
purpose  to  show,  as  the  German  perversion  of  the  struggle- 
and-survival  hypothesis  holds,  that  the  evolution  of  man 
has  consisted  largely  in  a  farther  differentiation  and  intensi- 
fication of  the  dominantly  brute  attributes,  with  an  infusion 
as  a  kind  of  by-product  from  the  struggle  for  existence,  of 
certain  "humanistic  sentimentalities,"  which  in  reality  are 
signs  of  weakness  and  must  be  suppressed.* 

And  this  perversion  by  German  science  and  philosophy  of 
Darwin's  teaching  is  rooted  very  deep  in  German  culture  and 
character.  The  straightforward,  common-sense  descriptions 
and  inductions  of  the  practical-minded,  country-dwelling, 
country-loving,  unacademic  English  naturalist  were  alto- 
gether too  simple  and  unsophisticated  to  satisfy  a  Kultur 
permeated  through  and  through  with  the  "highly  strained 
idealism"  of  Kant,  Fichte,  Hegel,  and  Schopenhauer.  The 
two  worst  errors  committed  by  Darwin  were  his  over-em- 
phasis on  the  natural  selection  hypothesis,  and  his  pro- 
pounding of  the  gemmule-pangenesis  hypothesis ;  and  it  is 
highly  characteristic  that  it  was  in  just  these  two  "strained" 
speculations  that  German  biology  and  practical  philosophy 
should  have  taken  up  Darwinism  the  most  ardently  and  over- 
worked it  the  most  absurdly  and  disastrously. 

My  examination  of  the  germplasm-determinant  theory  of 
Weismann  in  Part  I  of  this  book  has  revealed  something  of 
the  scope  and  nature  which  the  gemmule  fallac}^  was  destined 
to  assume  when  it  fell  subject  to  German  speculation.  The 
more  subtle  and  far-reaching  and  humanly  practical  conse- 
quences of  the  adoption  and  elaboration  of  the  struggle-and- 

*  The  effort  which  Dr.  George  Nasmyth  has  made  in  his  book  Social 
Progress  and  the  Darwinian  Theory  to  set  right  Darwin's  position  in 
this  matter,  ought  to  bear  fruit  after  a  while. 


Postscript  357 

survival  hy])()tlicsis  by  Griiiiaii  spcciilat  ion  lias  not  \(t  \nvi\ 
subjected  to  thor()ugli-«4()inn-  Ijiolo^lcal  criticism,  tliougli  sev- 
eral moves  in  tliis  direction  have  been  made. 

Even  the  realism  of  recent  (ierman  j)oHtii-al  and  economic 
tlieory  and  practice  is  a  "highly  strained"  sjK'culative  real- 
ism. This  philoso})hical  monstrosity  is  largely  attribntabh-, 
demonstrably  so  I  believe,  to  a  cultural  and  govirmnental 
system  in  which  the  principle  of  universal  organic  personality 
is  grossly  violated.  And  what  a  price  in  misery  and  blood 
and  treasure  the  whole  world,  but  old  Europe  ))articularly, 
is  paying  for  a  consummation  wliich  a  truer  ])hilos()])liy  of 
life  would  have  foreseen  and  forestalled ! 

Can  the  leaders  of  German  Kultur  be  convinced  of  the 
fundamental  fallacy  of  their  theory  of  human  and  nationjd 
life,  only  by  discovering  that  their  military  establishment, 
built  up  through  many  decades  of  ])atient,  costly  organiza- 
tion and  discipline,  but  under  guidance  of  a  ])hiloso})hy  of 
mechanism  and  brutism,  is  yet  incapable  of  overpowering  a 
military  establishment,  a  large  portion  of  which  may  be  im- 
provised in  the  course  of  a  few  months,  if  such  improvision 
be  under  guidance  of  a  philosophy  of  personality  and  hu- 
manism.'^ 


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. 


GLOSSARY 


AcROGEMALY.  A  discasc  charac- 
terized by  hypertrophy  of  the 
terminal  parts  of  the  body,  as 
of  the  face  and  extremities;  an 
outgrowth  involving  bony  and 
soft  parts. 

Adrenals.  A  pair  of  small  glands 
situated  in  front  of  the  kidneys. 
They  are  glands  of  internal  se- 
cretion, their  secretion  exercising 
in  particular  a  regulating  eflfect 
on  the  nerves  of  the  heart  and 
blood  vessels. 

Adrenin.  The  "active  principle" 
in  the  secretion  of  the  adrenal 
glands. 

Ai.cYoxARiA,  include  many  corals 
and  other  coelenterate  animals 
with  eight  mesenteries  and  eight 
tentacles. 

Al\t:olar.  In  anatomy,  a  numer- 
ously-pocketed, or  sacculated, 
structure,  typified  by  the  ter- 
minal cavities  of  the  lungs,  but 
occurring  in  various  tissues;  be- 
lieved to  constitute  also  one  kind 
of  protoplasmic  structure. 

Amoeba.  A  unicellular  animal,  a 
genus   of  rhizopodous   Protozoa. 

Ampiiibiax.  An  animal  living 
both  in  water  and  on  land. 
Properly  a  class  of  vertebrates 
whose  young  are  typically  aquat- 
ic and  respire  by  gills;  examples, 
frogs,  toads  and  salamanders. 

\:MPHioxrs,  literally  pointed,  or 
sharp  at  both  ends.  The  cur- 
rent name  for  one  of  the  very 
simplest  and  lowest  vertebrate 
animals  occurring  in  the  sand 
nnd  mud  of  the  seashore  in 
many  parts  of  the  world. 

AxABOLic,   the   chemical    up-build- 


ing of  the  living  body;  construc- 
tive metabolism. 

Antibodies.  "The  products  of  a 
reaction  of  the  body  towards  a 
natural  or  artificial  introduction 
into  it  of  certain  foreign  sul)- 
stances,  bacteria  and  their  poi- 
sons, vegetable  poisons  of  other 
kinds,  and  various  albuminoids." 
The  name  antil)odies  has  refer- 
ence to  the  antagonism  these 
products  have  for  the  introduced 
substances. 

Antigens.  Substances  the  reac- 
tion of  which  with  the  living 
body  produce  antibodies. 

AsciDiANS.  Marine  animals  hav- 
ing a  gelatinous  or  leathery  en- 
velope containing  cellulose.  In 
the  larval  stage  a  notochord  or 
forerunner  of  the  vertel)ral  col- 
umn is  present.  Some  free  liv- 
ing species  retain  the  notochord 
all  through  their  lives. 

Axon.  The  long,  slender,  S|)arce- 
ly-branched,  fibrillar  process  of 
a  ganglion  cell;  contrasted  with 
the  shorter,  more  branched,  more 
irregular  chndron. 

Axosri'LE.  \  s'endcr,  flexible  rod 
of  organic  substincc  forming  a 
supporting  axis  for  the  body  in 
some   Prntn/on. 

BioGKN.  I.iteriry  life  ])roducer. 
Imarinniy  i-ltimate  units  of  life. 
Such  special  si<Tnircnnrc  as  the 
"bingen  tlienry"  his  over  otlit-r 
theories  whi'h  mike  iniatrinary 
vital  or  pliysiilorical  units  a 
goal  of  the  ultinntc  cxplanntion 
of  Wfo,  is  foind  in  th«^  fact  thit 
the  I  ip'>Tn  theory  aims  to  be 
more     dclnitelv     chemical     tlian 


377 


378 


Glossary 


the  others.  The  German  physi- 
ologist Max  Verworm  has  elab- 
orated this  speculation  more 
fully  than  has  any  one  else. 

BioPHOR.  Literally  life  carrier. 
Biophors,  the  imaginary  ultimate 
vital  units  of  the  Weismannian 
system  of  speculative  biology, 
differ  from  biogens  in  the  fact 
that  Weismann,  not  being  a 
chemist  or  even  a  physiologist, 
but  a  zoologist  interested  in  re- 
production and  heredity,  rather 
than  in  function  generally,  did 
not  undertake  to  put  his  specu- 
lation on  a  chemical  basis. 

Bioplasm.  Formative  living  mat- 
ter; not  differing  in  any  but  a 
speculative  way  from  proto- 
plasm. 

BioTic.  Pertaining  to  living 
beings. 

Blastogexesis.  Reproduction  by 
budding,  as  used  in  this  book; 
in  general,  propagation  from  an 
undifferentiated  germinal  mass. 

Blastomeres.  The  first  segments 
or  cells  formed  by  the  division 
of  the  ovum. 

Blastozooids.  The  united  individ- 
uals produced  by  budding,  and 
constituting  the  colony,  or  cor- 
mus  in  the  compound  ascidians. 

Blastula.  The  stage  of  develop- 
ment of  the  embryo  from  the 
ovum  in  many  animals,  in  which 
the  organism  consists  of  a  hol- 
low sphere  the  wall  of  which  is 
composed  of  a  single  layer  of 
cells. 

Bryozoa.  Literally  "moss  ani- 
mals," from  the  resemblance, 
fancied  more  than  real,  of  some 
of  the  species  to  mosses;  also 
called  Polyzoa.  Marine  animals 
occurring  abundantly  on  all 
shores.  Most  of  the  species 
propagate  by  budding  as  well  as 
by  eggs  and  sperm,  the  bud-pro- 
duced individuals  remaining  at- 
tached to  one  another  to  form 
colonies,  as  in  many  hydroids  and 


ascidians.  Each  individual  con- 
sists of  a  body  proper  bearing 
a  circle  of  tentacles,  and  an  en- 
veloping case  often  calcareous, 
into  which  the  body  may  be 
quickly  and  completely  re- 
tracted. 

Calymma.  The  much-vacuolated 
portion  of  the  body  of  radio- 
laria,  situated  outside  the  central 
capsule,  the  vacuoles  containing 
fluid  impregnated  with  gas.  The 
main  office  of  the  structure 
seems  to  be  in  connection  with 
the  flotation  of  the  animals. 

Cambium.  The  layer  in  woody 
plants  between  the  outer  dead 
layer,  or  bark,  and  the  inner 
dead  mass,  or  wood  proper,  from 
which  new  tissue  is  formed;  the 
true  growing  part  of  plants 
which  live  several  years  and  at- 
tain a  large  size. 

Cartesian  Philosophy.  The  mode 
of  viewing  man  and  nature  in- 
augurated in  modern  times  by 
Rene  Descartes.  The  most  dis- 
tinctive thing  about  it  is  the 
sharpness  with  which  the  dual- 
ism, or  antithesis,  between  mind 
and  matter  stands  out  in  it.  Its 
great  practical  importance  for 
the  present  era  lies  in  its  genetic 
relationship  to  psycho-physical 
parallelism  in  psychology,  and  to 
all  forms  of  idealism  in  philos- 
ophy. 

Cektrolecithal  Eggs.  Eggs  in 
which  the  protoplasmic  portion 
and  nucleus  constitute  a  surface 
layer,  the  inner  mass  being 
chiefly  yolk,  that  is  food  ma- 
terial for  the  future  embryo. 
The  eggs  of  most  insects  and 
crustaceans  are  of  this  type. 

Cephaltze.  The  tendency  among 
animals  for  a  head  to  become 
differentiated  from  the  rest  of 
the  body. 

Chemical  Messengers.  Substances 
produced  by  the  organism, 
either     in     special     glands,     the 


Glossary 


379 


glands  of  internal  secretion,  or 
in  general  tissues,  and  carried 
by  the  blood  and  1}  niph  over  the 
body  generally,  to  influence  the 
growth  or  functioning  of  other 
tissues.  Horviones  is  another 
name  given  to  these  sul)stances. 

CiiEMOTOOPic.  Pertaining  to  the 
reaction  of  organisms  to  chem- 
ical stimuli. 

CiiROMATix.  The  finely  granular 
substance  most  distinctive  of  the 
cell-nucleus.  Its  name  conies 
from  the  readiness  with  which 
it  is  colored  by  many  dye-stuffs. 

CiiROMATOPHOREs.  Pigment-bear- 
ing sacs,  often  single  cells,  in 
plants  and  animals.  It  is  by 
means  of  these  that  the  rapid 
color  changes  in  the  skin  of 
nianv  ap.imals  are  accomplished. 

C'iiROMiDoso3iEs.  One  of  the  many 
names  given  to  minute  specially 
stainable  bodies  in  the  cytoplasm 
of  many  cells. 

Chromosomes.  The  more  or  less 
definite  bodies  of  the  cell-nu- 
cleus into  which  the  chromatin 
granules  are  grouped.  Their 
constancy  of  structure  and  rela- 
tion to  hereditary  characters 
have  given  them  great  promi- 
nence in  much  of  recent  biolog- 
ical theory. 

Chromopiiil.  Any  body  in  the  liv- 
ing organism  that  has  an  avidity 
for  staining  reagents. 

Coccus.  In  the  classification  of 
bacteria  according  to  their 
shapes,  those  which  are  spherical 
are  called   cocci. 

CoELEN'raRATA.  A  group  of  ani- 
mals that  have  a  digestive  cav- 
ity, but  nothing  corresponding 
to  the  abdominal  cavity;  also 
called  radiata  from  the  radial 
arrangement  of  the  body. 

Cortical.  Pertaining  to  the  cor- 
tex, or  outer  layer  of  an  organ, 
as  of  the  brain,  l)ark  of  tiie  tree, 
&c. 

Cretin.       An    individual    affected 


with  cretinous  disease,  a  disease 
characteri/ed  by  certain  bodily 
deformities  aiul  mental  inijiair- 
nicnts.  The  malady  frtfjiii-ntly 
accompanies  goitre,  and  is  now 
considered  due  to  deficiency  in 
secretion  of  the  thyroid  gland. 

CuiSTACEA.  A  class  of  invt-rte- 
l)rate  animals  lielonging  to  the 
great  ph\luin  arthroj)oda,  briefly 
characterized  by  their  exoskele- 
ton  and  paired  jointed  aj)i)en- 
dages. 

Cytology.     Science  of  the  cell. 

CYTOPLAs:Nr.  Substance  of  the  cell- 
body,  as  opposed  to  the  cell-nu- 
cleus. 

Determixants.  That  particular 
class  of  imaginary  ultimate  vital 
units  by  which  the  develo])incnt 
of  hereditary  attributes  is  deter- 
mined. They  were  invented  by 
Weismann,  and  were  conceived 
to  constitute  the  germ-plasm, 
and  to  be  located  primarily  in 
the  chromosomes  of  the  egg  and 
sperm.  In  later  speculation  de- 
terminer is  used  more  frccjuently 
than  determina/f/ — on  some  ac- 
count  that   is   not  clear. 

Diatoms.  An  immense  group  of 
aquatic,  unicellular  alga-  espe- 
cially characterized  liy  their 
firm,  box-like,  regularly-sliaj>ed, 
chitinous  shell. 

Dixoflagei.lates.  Literally  or- 
ganisms which  are  two-lashed, 
owing  to  the  two  flagella  j)os- 
sessed  by  most  of  the  species. 
A  group  of  a(juatic  uniei-llnlar 
organisms  almost  as  nunu-rous  as 
the  diatoms.  The  photo-syiithe- 
siziiig  power  of  living  substance 
j)Ossessed  by  these  two  groups, 
and  their  enormous  abundance 
at  and  near  the  siirface  of  the 
bodies  of  water  in  which  they 
live,  make  tlwin  fundamentally 
important  for  all  the  life  of  the 
waters   of   the   earth. 

DiSTAi.  A  connuon  anatomical 
term     signifying    away     from    a 


880 


Glossary 


given  point,  usually  some  defi- 
nite feature,  as  the  attachment 
of  a  muscle,  taken  as  a  point  of 
reference. 

DoKSo- Ventral.  A  term  much 
used  in  the  anatomy  of  the 
higher  animals  to  signify  a  di- 
rection from-back-to-belly  of  a 
creature. 

DuoDEXu^r.  The  first  portion  of 
the  small  intestine  between  the 
stomach  and   the  Jejunum. 

Dynamic  Center.  A  phrase  used 
rather  freqiently  in  recent  biol- 
ogy, especially  in  the  biology  of 
the  cell,  to  express  the  concep- 
tion that  certain  structures,  as 
the  centrnsome,  nre  in  someway 
not  clearly  sncifiab'e  the  "seat" 
of  various  vital  activities.  The 
phrase  has  some  such  implica- 
tion for  general  ])hysi'ilogy  as 
"nerve  center"  had  and  with 
many  still  has,  for  nerve  phys- 
iology. 

Ecological.  Pertaln.'na:  to  Ecol- 
ogy, the  science  of  organisms  in 
relation  to  their  mtural  environ- 
ments. This  old  but  newly  ap- 
preciated and  named  branch  of 
the  science  of  living  nature,  may 
properlv  h,e  regarded  as  the  nat- 
ural history  of  plants  and  ani- 
mals modified  to  meet  the  mod- 
ern demands  of  comprehensive- 
ness and  exactness  in  dealing 
with  a  great  province  of  nat- 
ural phenomena. 

Ectoplasm.  The  outermost,  some- 
what denser  layer  of  protoplasm 
in  many  cells,  especially  in  many 
unicellular  animals.  Opposed  to 
endoplasm,  the  inner,  more  fluid 
mass.  The  ectoplasm  in  proto- 
zoans corresponds  to  the  skin  of 
higher  animals.  The  presence  of 
a  more  or  less  sharply  set-off 
outer  layer  or  membrane  or  skin 
in  all  organisms  whatever  is 
coming  to  be  recognized  as  hav- 
ing a  more  fundamental  physio- 
logical meaning  than  that   of  a 


protection  for  the  delicate  parts 
underneath,  now  that  so  much  is 
being  learned  by  physical  chem- 
istry  about   surface   phenomena. 

Energy.  Work  and  capacity  to  do 
work.  It  is  important  to  note 
that  work  and  capacity  to  work 
necessarily  imply  some  object, 
organic  or  inorganic,  to  do  the 
work,  and  hence  that  when  the 
energy  of  a  horse,  or  of  a  stream 
of  water,  is  s})oken  of  the  word 
energy  has  a  very  different 
meaning  from  what  it  has  in 
such  a  phrase  as  the  "energy 
conception"  of  nature  or  of  the 
organism,  the  implication  in 
these  cases  usually  being  that 
energy  is  the  real  essence  of 
nature  and  of  the  organism,  the 
shape  and  other  so-called  static 
attributes  which  all  bodies  pre- 
sent, being  only  incidental  and 
mere   appearances. 

Enzyme.  A  chemical  substance 
produced  by  an  organism,  plant 
or  animal,  to  the  end  of  bring- 
ing about  chemical  transforma- 
tion in  other  substances,  but 
without  itself  being  transformed. 
The  ptyalin  of  saliva  by  which 
starch  is  changed  into  sugar,  is 
typical.  Enzymes  play  a  very 
great  part,  especially  in  diges- 
tion and  nutrition,  in  the  physio- 
logical processes  of  all  organ- 
isms. 

Epige nests.  That  theory  of  devel- 
opment of  the  individual  organ- 
ism which  holds  the  organs  and 
parts  to  be  actually  new  produc- 
tions, and  not  merely  enlarge- 
ments or  actualizations  of  wihat 
already  existed,  this  latter  con- 
ception of  development  consti- 
tuting the  theory  of  preforma- 
tion. Although  these  opposing 
theories  were  debated  with  fury, 
almost,  some  years  ago,  little  is 
heard  about  them  now  though 
none  of  the  particular  problems 
around     which     the     discussions 


Glossary 


381 


centered  can  be  said  to  have 
been   solved. 

Epimorpiiosis.  The  mode  of  re- 
generation of  organisms  in 
which  a  multiplication  of  cells 
on  the  surface  of  injury  is  first 
produced,  then  from  tliis  ''em- 
bryonal tissue,"  the  new  organ 
or  part  is  formed;  contrasting 
with  morphahwis,  a  mode  of 
new  formation  which  consists  in 
a  direct  transformation  of  an 
already  existing  part  into  the 
new   part. 

Factor  (in  Genetics).  A  hypo- 
thetical unit  of  structure  or  of 
chemical  composition,  containeJ 
in  the  germ-cell,  which  in  some 
way  is  held  to  condition  the  de- 
velopment of  a  particuhir  char- 
acter in  the  adult,  or  of  a  com- 
plex of  characters  which  are 
transmitted  in  constant  associa- 
tion with  one  another.  Factors 
are  believed  to  interact  with  one 
another  in  development,  and  at 
times  to  be  so  "linked"  that  they 
are  only  partially  independent  in 
transmission. 

Flagellum.  a  lash-like  appen- 
dage or  large  cilium  serving  as 
an  organ  of  locomotion  in  some 
Protozoa  and  some  bacteria. 

FoRAMTXiFERA.  A  class  of  rhizo- 
podous  marine  Protozoans,  usu- 
ally  having  a  porous  shell. 

Formative  Stuffs.  Hypothetical 
substances  which  are  supposed 
to  be  formed  in  one  part  of  an 
organism  and  transported  to  an- 
otiier  part,  there  to  produce,  or 
to  influence  the  production  of 
new  organs.  For  example,  sev- 
eral botanists  have  supposed  that 
the  flower  substance  of  some 
plants  is  actually  produced  in 
the  leaves. 

Gamete.  A  reproductive  cell 
which  unites  with  another  repro- 
ductive cell  to  form  a  zygote. 

Gastrula.  That  stage  of  embry- 
onic  development   in   many   ani- 


mals which  consists  of  two  germ- 
layers  inclosing  a  central  cavity. 
It  is  produced  from  the  blastuli 
(which  see)  by  liic  in-sinking  of 
one-half  of  this  into  the  other. 

Gemmiparois.  Producing  pcnnnu', 
or  buds  (reproducing  by  bud- 
ding), aiiplical)le  to  botli  plants 
and    many    animals. 

Gemmule.  In  the  origind  anJ 
]iroper  sense  a  small  aggregation 
of  cells  set  apart  in  the  tissues 
of  some  plants  aiul  animals, 
notably  in  many  sponges,  for  the 
puri)Ose  of  reproduction.  In  ori- 
gin and  structure  gcmnnilcs  arc 
more  like  brds  than  eggs, 
thouf.'h  the  end  served  is  very 
similar  to  tliat  st-rved  by  sercU. 
In  a  secondary  and  wlinlly  hyjio- 
theti^al  sense,  gennnules  are 
imaginary,  minute  bodies  given 
off  by  ail  the  tissue  ceils  of  an 
organism  and  assembled  in  the 
germ  cells,  there  to  cause  the 
development  of  the  next  genera- 
tion. This  taking  of  a  very  con- 
crete name  from  botany  and 
zoology,  and  using  it  in  a  wliolly 
imaginary  way  to  explain  hered- 
itary development  was  due  orig- 
inally to  Charles  Darwin,  but 
with  more  or  less  unimportant 
variations  of  meaning  has  since 
iieen  resorted  to  by  many  of  the 
best  known  biologists. 
This  exami)le  indicates  the  great 
importance  for  biolog)-,  espe- 
cially for  the  biology  of  rrpro- 
duction  aiul  develo|>ment,  of  dis- 
tinguishing betwct-n  the  same 
terms  used  in  a  strictly  objective 
and  descrij^tive  sense  on  the  (»ne 
hand,  and  in  hypothetical,  or 
pure!v  imairinar\'  sense  on  the 
other. 

Gene.  A  term  much  used  in  prrs- 
ent-dnv  gen<>tic.il  science,  but  ap- 
|>arenlly  not  differing  in  any  sig- 
nificiuit  particular  from  factor 
(which  see). 

Genetic.      Pertaining    to    gt-mtics. 


■382 


Glossary 


evolutionary  science  dealing  with 
natural  propagation  and  devel- 
opment, the  interest  centering  at 
present  in  that  portion  of  devel- 
opment which  is  hereditary  and 
involves  sex  cells. 

Germ-plasm.  Actually  all  the  pro- 
toplasm of  the  germ-cells  which 
participates  in  development;  the- 
oretically merely  the  small  por- 
tion of  the  germ-cells  supposed 
to  be  "hereditary  substance." 

GoxAD,  A  mass  of  undifferen- 
tiated, generative  tissue  from 
which  the  male  and  female  re- 
productive glands  originate. 

GoxoPHORE.  The  ultimate  gener- 
ative zooid  of  a  hydrozoan,  giv- 
ing origin  directly  to  the  genera- 
tive elements. 

Hectocotylized.  Applied  to  the 
remarkably  altered  condition  as- 
sumed by  one  of  the  arms  of  the 
male  cephalopod  to  make  it  an 
organ  for  impregnating  the  fe- 
male. 

Heliotropic.  Responding  to  the 
stimulus  of  sunlight. 

Heteromorphosis.  a  kind  of  re- 
generation in  which  the  part  pro- 
duced is  different  from  that 
which  was  lost,  as,  for  example, 
when  an  antenna-like  structure 
grows  in  the  place  of  an  eye- 
stalk,  in  some  crustaceans  when 
the  eye  stalk  is  cut  off. 

Histogenesis.  The  process  of  tis- 
sue genesis,  or  production,  from 
undifferentiated  cell  masses,  in 
plants  and  animals. 

Histology.  The  science  of  tissues, 
plant  or  animal;  microscopical 
anatomy. 

HoRMoxE.        Literally      something 

_  which  excites  or  stirs  up.  Orig- 
inally and  strictly  applied  to 
those  internal  secretions  (which 
see),  the  office  of  which  is  to 
incite  the  parts  on  which  they 
act  to  greater  activity.  But  in- 
ternal secretions  are  also  known 
now  which  retard  or  inhibit  the 


action  of  the  part  they  affect; 
and  to  these  it  has  been  proposed 
to  apply  the  term  chalone,  that 
which  slackens.  But  some  phys- 
iologists use  hormones  as  syn- 
onymous with  "internal  secre- 
tions." 

Homonymous.  As  used  in  this 
book,  an  anatomical  term  refer- 
ring to  the  different  members  of 
a  series  which  differ  more  or 
less,  but  still  all  have  the  same 
general  name.  Thus  all  the 
pairs  of  appendages  of  a  lobster 
are  homonymous,  or  ambulatory 
appendages  originally,  although 
used  for  a  variety  of  purposes 
now. 

Hydranth.  One  of  the  bud-pro- 
duced polyps  of  a  hydroid  col- 
ony. 

Idioplasm.  Literally  plasm  which 
is  very  specially  one's  own.  First 
used  to  designate  the  hypothet- 
ical part  of  the  germ-cells  which 
is  supposed  to  be  alone  respon- 
sible for  hereditary  transmission. 
Idioplasm  may  be  regarded  as 
the  historical  antecedent  of 
germ-plasm   (which  see). 

Interstitial.  Pertaining  to  or  sit- 
uated in  an  intervening  space;  a 
term  much  used  in  anatomy  to 
signify  within  an  organ. 

Internal  Secretion.  The  term 
has  long  been  used  in  the  phys- 
iology of  the  higher  animals,"  in 
contradistinction  to  "external  se- 
cretion," to  designate  the  prod- 
ucts of  glands,  like  the  thyroid, 
which  discharge  their  products 
into  the  blood  or  lymph,  instead 
of  upon  the  surface  of  the  body 
or  into  the  digestive  or  some 
other  cavity  of  the  body.  The 
existence  of  internal  secretions 
was  known  long  before  anything 
was  known  about  their  use; 
hence  this  non-committal  name, 
so  far  as  function  is  concerned. 
The  recent  discovery  of  their 
office    has    suggested    the    name 


Glossary 


383 


hormone  (which  see)  for  theiu, 
and  has  revealed  their  great  im- 
portance not  only  for  physiology, 
hut  for  philosophical  Inology. 

Involution.  Literally  inrolling,  or 
inwrapping.  In  descriptive  hiol- 
ogy  used  to  signify  the  return  of 
an  organ  to  its  original  or  nor- 
mal condition  after  some  violent 
or  pronounced  deformation  of  it. 
Sometimes,  but  apparently  un- 
justifiably, used  as  a  synonym  of 
degeneration.  Since  the  doctrine 
of  evolution  has  become  prom- 
inent in  biology,  a  process  the 
oi)])osite  of  evolution  has  l)een 
thought  by  some  to  be  necessary, 
and  to  this  involution  has  been 
applied. 

Jei.ly-fish.  In  the  interest  of  dis- 
criminative knowledge,  the  habit, 
rather  common  among  people 
who  have  the  opportunity  to  see 
the  transparent,  somewhat  gela- 
tinous-appearing animals  of  the 
ocean,  of  calling  them  all  "jell}'- 
fishes"  should  be  abandoned.  The 
name  should  be  restricted  to  the 
regularly  disc-  or  dome-shaped, 
tentaculated  animals  belonging 
to  the  coelenterate  phylum,  thus 
enlarging  the  bounds  of  defmite, 
popular  zoological  information, 
by  recognizing  that  marine  ani- 
mals of  several  large  and  very 
distinct  classes  have  this  general 
consistency  and  appearance. 

Karyoplas^i.  a  cytological  name 
referring  to  the  substance  or 
plasm  distinctive  of  the  cell-nu- 
cleus. 

Karyosome.  a  small,  discrete, 
rather  constant  body  which 
stains  readily,  contained  in  the 
cell-nucleus;  frequently  synony- 
mous with  nucleolus. 

Katabolism.  The  down-breaking, 
or  descensive  phase  of  metabol- 
ism; the  opposite  of  amibolism 
(which  see). 

Ktneto-nuci.eus.  One  of  the  nu- 
clei  in  the  two-nuclear  protozoa 


supposed  to  be  concerned  in 
some  special  way  with  the  move- 
ment of  the  flagclla  or  cilia  of 
tliese   animals. 

l.A.-\iKi,r.AE,  singular  lavtella.  A 
term  much  used  in  anatomy  to 
designate  the  thin  plates,  .scales, 
etc.,  that  are  so  nuiiu-rous  and 
varied  in  form  and  size  in  nearlv 
all  organisms. 

Larva.  Properly  apj)Iied  only  to 
stages  in  the  lives  of  iiulividii.il 
animals  which  ])ass  into  succeed- 
ing stages  through  ;i  deep-sealed 
nu'tam()rj)hosis,  as  for  example 
the  grub  or  maggot  of  a  fly,  and 
its  transformation  into  the  adult. 
Larval  stages  aiul  ])rofoiin(l 
metamorphoses  are  very  con)tnon 
and  widespread  in  the  animal 
kingdom. 

LiMULus.  The  technical  genus 
name  for  the  horse-shoe  crab,  an 
animal  of  special  interest  to  gen- 
eral zoology  in  several  ways. 

Macronuci.evs.  In  the  infusoria,  a 
group  of  jirotozoans,  there  is  one 
large  nucleus  and  one  or  several 
nnich  smaller  nuclei.  'I'he  first  is 
called,  from  its  relatively  large 
size,  the  ;/K»r/o-nucleus;  the 
others  m/r-ro-iuu-Iei.  From  th<- 
behavior  of  the  two  kiiuls  of  nu- 
clei at  conjugation  and  (livi>ioM, 
the  micronuclei  are  known  to  be 
intimately  comiected  with  these 
processes,  while  the  macromicleus 
seems  to  be  more  coneeriu-tl  with 
the  nutritive  functions  of  the 
arn'nial. 

IMANiiuur.-M.  In  n)orj)hoIogv  n 
part  or  organ  which  resembles  a 
handle;  specially  the  clajipcr- 
like,  or  handle-like  portion  of  a 
medusa  which  is  found  within 
the  "bell."  The  animal's  mouth 
is  at  the  erul  of  i\\e  manubrium, 
aiul  most  of  its  digestive  cavity 
within  the  stalk  of  the  maiui- 
iirium. 

•ALvniix.  In  biology  the  ground 
substanie  in  which  cells  arc  em- 


384 


Glossary 


bedded  in  some  tissues,  and 
which  is  produced  as  a  secretion 
by  the  cells.  It  is  one  kind  of 
intercellular  substance.  The 
opalescent,  almost  homogeneous 
chief  mass  of  ordinary  cartilage 
is  a  typical  matrix. 

Melaxin.  a  rather  general  term 
in  biology,  especially  in  zoology, 
applied  to  dark  brown  to  black 
pigments. 

Merotomy.  The  automatic  cutting 
off  of  jDarts  or  segments  in  liv- 
ing organisms. 

Mesenchyme.  Undifferentiated 

mesoderm  that  produces  con- 
nective tissues,  some  muscles, 
and  certain  other  structures  in 
the  animal  body. 

Metabolic.  Pertaining  to  metab- 
olism, the  process  of  chemical 
building  up  and  breaking  down 
in  the  living  organism. 

Metameric.  Pertaining  to  the 
longitudinal  series  of  parts  or 
joints  into  which  the  bodies  of 
many  higher  animals,  such  as 
earthworms,  lobsters  and  fishes, 
are  divided. 

Metaplastic.  Pertaining  to  meta- 
plasm — applied  to  changes  which 
cells  sometimes  undergo  from 
one  plasmic  type  to  another;  also 
applied  to  certain  supposedly 
lifeless  inclusions  in  the  proto- 
plasm of  cells. 

Metazoa.     Multicellular  animals. 

MiCROxucLEus.     See  macronucleus. 

Microphyle.  In  botany  and  zool- 
ogy the  aperture  in  the  coats  of 
the  ovule  and  ovum  through 
which  the  male  fertilizing  cell 
penetrates. 

MoKERA.  Hypothetical  simple 
structureless  masses  of  proto- 
plasm (without  any  nucleus). 
Assumed  by  Haeckel  as  the  low- 
est members  of  the  evolutionary 
series.  Advance  of  knowledge 
has  found  no  evidence  of  such 
organisms. 

MoRPHALLAXis.     A  kind  of  regen- 


eration in  which  part  of  an  or- 
ganism transforms  directly  into 
a  new  and  different  part. 

Morphological.  Pertaining  to 
morphology,  the  science  of  form 
and  structure. 

Morula.  A  stage  in  the  embryonic 
development  of  many  animals,  in 
which  the  ovum  has  completely 
segmented,  but  the  segmentation 
cavity  has  not  yet  been  formed. 

Myoneme.  a  thread-like  contrac- 
tile structure  in  the  cytoplasm  of 
certain  higher  protozoa. 

Nematophore.  a  body  of  defense 
and  offense  developed  in  certain 
hydroids,  consisting  of  a  chitin- 
ous  receptacle  in  which  thread- 
cells  are  immersed;  the  nettling 
organs  on  the  tentacles  of  large 
jelly-fishes. 

Neural.     Pertaining  to  nerves. 

Neuroblasts.  Undeveloped  nerve 
cells. 

NucLEo-PLASM.  Nuclcar  substance, 
including  the  different  nuclear 
ingredients. 

NucLEo-PROTEix.  Ouc  of  the  com- 
pounds of  nucleins  and  paranu- 
cleins. 

CEdema.  Dropsy,  a  vasomotor 
neurosis  characterized  by  non- 
inflammatory swellings  on  vari- 
ous parts  of  the  body. 

Ontogexy.  The  development  of  an 
individual  organism  from  germ 
to  completed  or  adult  stage. 

Organelle.  A  little  organ,  and 
organoid,  organ-like,  are  terms 
applied  to  the  organs  of  unicel- 
lular plants  and  animals,  not  so 
much  because  of  their  small  size 
and  indefiniteness  of  form  and 
structure  as  on  account  of  the 
theory  that  a  true  organ  must 
be  composed  of  cells,  and  cannot 
be  a  part  of  a  cell.  These  terms 
are  among  the  sequelae  of  the 
cell-theory. 

Orienting.  Finding  or  fixing  the 
positions  or  directions. 

Pangen,   and   Pangenesis.     These 


Glossary 


385 


terms,  basal  in  Darwin's  famous 
hypothesis  of  heredity,  mean  all- 
generator  and  all-generative  only 
in  the  sense  that  all  parts  of  the 
body  of  the  organism  give  off 
gemmules  (which  see),  which 
assemble  in  the  germ-cells  to  en- 
able these  to  reproduce  the  or- 
ganism. Thus  the  -pan,  or  all- 
gencrative  power  was  conceived 
as  having  its  original  "scat"  in 
the  organism  all-in-all.  In  other 
words  Darwin's  speculation  was 
almost  diametrically  opposed  to 
the  transformation  it  has  under- 
gone latterly,  especially  in  the 
prolific  mind  of  Weismann,  the 
germ  cells  alone,  or  rather  the 
germ-plasm  being  the  all-genera- 
tor, according  to  these  specula- 
tions. 

Parathyroids.  Small  glands  lying 
near  the  thyroid  but  not  func- 
tionally connected  with  the  lat- 
ter. 

Parthexogexesis.  Reproduction 
by  means  of  unfertilized  eggs. 

Pellicula.  The  cuticle  or  outer- 
most body  membrane  in  some 
unicellular  and  other  low  organ- 
isms. 

Phase.  This  old  and  familiar  word 
has  taken  on  new  and  greater 
importance,  both  scientific  and 
philosophic,  with  the  recent  ad- 
vance of  knowledge  in  the  region 
of  over-lap  between  physical  and 
chemical  phenomena,  this  ad- 
vance making  what  is  generally 
called  physical  chemistry.  A 
'phase  in  pure  physics,  as  it  may 
be  called,  has  reference  to  the 
position  of  the  particles  of  a 
body  when  the  particles  are  un- 
dergoing change.  For  example, 
corresponding  particles  in  two 
succeeding  waves  of  water  or  air 
are  in  the  same  phase.  In  phys- 
ical chemistry  phase  has  refer- 
ence not  to  position  but  to  state 
or  condition  of  the  constituent 
particles  of  a  heterogeneous,  or 


unlike  system.  Thus,  a  combina- 
tion of  liijuid  water  (in  comiiioii 
language  just  water)  and  .vilid 
water,  or  ice,  is  a  two-phase  sys- 
tem of  water.  I'ljilosophirally 
viewed,  the  great  signifieariee  of 
phases  is  that  the  j)ositions  aiul 
statesof  the  particles  are  |)ossilii«-, 
even  conceivable,  only  in  relation 
to  the  larger,  containing  part  or 
whole.  Something  of  the  ln-ar- 
ing  of  this  on  the  theory  of  plu- 
ralism (which  see),  when  this 
theory  is  apj)roaelied  from  IIm 
strictly  objective  side,  will  1..- 
easily  seen. 

Phlogiston'.  An  imaginary  sui»- 
stance  formerly  supjjoscd  to  ex- 
ist in  all  combustible  boilies,  and 
to  be  the  cause  of  fire  and  flame. 
For  nearly  a  century  before  the 
discovery  of  oxydation  as  the 
true  cause  of  fire,  by  Lavoisier. 
the  phlogistic  theory  dominated 
much  of  chemical  science.  The 
chief  interest  in  the  theor}-  now 
is  in  its  relation  to  the  observa- 
tional and  logical  processes  in- 
volved in  interpreting  the  gen- 
erative processes  of  nature 
everywhere.  The  phlogistic  the- 
ory may  be  taken  as  a  type  of 
elementallstic  causal  cxj)lanation 
of   natural    jiroduction. 

Phvlogexic.  Pertaining  to  phy- 
logeny,  the  developnunt  of  the 
race;  concerning  ancestral  or- 
ganisms, real  and  hypoth«tieal. 

PiTuiTAHv  Cii.ANi).  A  glaiul  of  in- 
ternal secretion,  situated  at  the 
base  of  the  brain,  and  comuvted 
in  the  embryo  with  the  roof  of 
the  mouth. 

Pi.raALiSM,  philosoj)hicnl  (so  usctl 
in  this  book).  The  conception 
that  in  its  deepest  nature  tin* 
universe  is  multiform  anil  com- 
j)lex;  the  opposite  of  Moni*tn, 
the  conception  that  .some  .single 
J"'ssenee  or  Substance,  more  or 
less  known  or  tinknown,  is  tin* 
foundation  of  nil   tilings. 


386 


Glossary 


Pluteus.  Name  given  the  charac- 
teristic process-bearing  larva  of 
sea-urchins  and  their  near  rela- 
tives. These  larvae  are  of  con- 
siderable general  interest  because 
of  the  extensive  use  made  of  sea- 
urchin  eggs  in  experimental  em- 
bryology, the  eggs  being  easily 
olitained  and  easily  kept  in  the 
laboratory. 

Proteixs.  Nitrogenous  substances 
found  in  the  bodies  of  plants 
and  animals.  These  substances 
are  usually  considered  to  be  the 
most  fundamental,  from  the 
chemical  standpoint,  in  organic 
beings. 

Protista.  A  group  name  intended 
to  include  all  unicellular  organ- 
isms; i.e.,  both  protophyta,  one- 
celled  plants,  and  protozoa,  or 
one-celled   animals. 

PsEUDOPODiA.  Literally  false  feet. 
They  are  temporary  protrusions 
of  the  protoplasm  of  some  pro- 
tozoa, especially  of  the  rhizopo- 
dous  class,  typified  by  the  amoe- 
ba, the  name  having  reference  to 
the  locomotor  office  of  the  proc- 
esses. But  their  food-taking  and 
digesting  office  should  be  noted 
also. 

Ptyalik.  The  unorganized  fer- 
ment, or  enzyme  of  saliva,  chiefly 
instrumental  in  the  conversion  of 
starch  into  sugar. 

Radiolaria.  One  of  the  main  sub- 
divisions of  the  protozoa,  espe- 
cially characterized  by  their  gen- 
erally spherical  outline,  and  ra- 
diating structures,  some  soft  and 
extensile,  others  stiff  and  per- 
manent. The  radiolaria  are  al- 
most all  marine. 

Regulation.  Much  used  in  studies 
in  the  regeneration  of  organisms, 
to  express  the  power  many 
plants  and  animals  have  of  un- 

.  dergoing  structural  or  func- 
tional readjustments  in  order  to 
retain,  or  to  regain,  their  typical 
form;  a  significant  adaptation  of 


a    general    term    to    a    technical 
end. 

Reticular.  Net-like,  a  term  much 
used  in  anatomy,  as  many  por- 
tions in  both  plants  and  animals 
of  many  grades,  present  this 
type  of  structure,  though  the 
netting  never  has  the  regularity 
of  manufactured  netting. 

Rhizopoda.  The  great  subdivision 
of  the  protozoa  especially  char- 
acterized by  sending  out  pseudo- 
podia  (which  see).  Amceba  is 
usually  mentioned  as  the  type  of 
this  subdivision,  but  the  larger 
number,  probably,  of  rhizopods 
possess  shells  of  one  sort  and  an- 
other, while  amoeba  is  entirely 
naked   during  all  its  active  life. 

Sarcode.  Literally  like  flesh.  The 
name  originally  applied  to  what, 
under  microscopic  examination, 
seemed  to  be  the  fundamental 
living-substance  of  animals.  La- 
ter discovered  to  correspond  to 
what  was  known  as  protoplasm 
in  the  cells  of  plants. 

Sarcodyctium.  a  protoplasmic 
network  of  the  surface  of  the 
calymma  of  a  radiolarian. 

Self-differentiatiok,  Self-regu- 
LATiox,  &c.  It  is  not  without 
philosophical  significance  that 
the  term  self  has  forced  its  way 
into  technical  biology,  something 
as  it  has  into  technical  philoso- 
phy. In  biology  the  term  is  par- 
ticularly common  in  connection 
with  developmental  phenomena 
and  has  reference  to  operations 
which  depend  primarily  on  the 
organism  itself,  and  can  be  re- 
ferred to  "external  factors"  only 
remotely  and  in  a  round-about 
way. 

Sericteries.  Glands  by  which  silk 
and  silk-like  substances  are  se- 
creted in  many  insects. 

Soma,  Somatic.  The  body  and 
pertaining  to  the  body.  Much 
used  in  later  discussions  of 
heredity  in  a  strongly  hypothet- 


Gloss  nr/j 


88' 


ical  sense,  to  indicate  the  com- 
plete in(le))en(lence,  so  far  as 
development  is  conccriu'd,  of  the 
body  from  the  germ.  Tlic  an- 
tithesis is  often  made  stronger  by 
sjx'aking  of  the  snhstance  of  the 
body  and  the  substance  of  the 
gern),  using  the  terms  soinaflr- 
phisin  and  germpUism.  From 
the  ])hilnsophical  standpoint  it  is 
instructive  to  comj)are  the  the- 
oretically complete  separation  of 
body  and  germ  in  modern  gen- 
etics, with  the  theoretically  com- 
plete sejiaration  of  body  and 
soul  in  philosophy  and  psychol- 
ogy. 

SpFx'iFiciTi'.  The  state  of  being 
specific,  that  is,  of  being  mani- 
fested as  ))henomena  distinguish- 
able from  all  other  ])henomena. 
The  grou])  of  terms  kindred  to 
.specific  and  .species,  long  import- 
ant in  systematic  and  taxonomic 
biology,  are  becoming  increas- 
ingly so  with  the  advance  of 
knowledge,  <'S)H'cially  in  tlie  do- 
mains of  the  chemistry  of  differ- 
ent kinds  of  organisms,  and  of 
coni})arative  behavior  and  psy- 
chology. 

Sporazoa.  One  of  the  main  sub- 
divisions of  the  protozoa  a  lead- 
ing characteristic  of  which  is  in- 
dicated by  the  name,  that  char- 
acteristic being  the  commonness 
with  which  propagation  occurs  in 
the  group  by  means  of  spores 
produced  within  the  body  of  the 
animal.  By  far  the  greater  num- 
ber of.  the  species  of  the  grou]) 
are  jiarasitic,  many  of  them  dis- 
ease producing. 

SFourr.ATioN.  The  process  of  con- 
verting into  spores,  as  in  the 
s])ora/,oa,  in  some  other  animals, 
and  in  many  )>lants.  Spores  dif- 
fer from  eggs,  on  the  one  h.ind, 
and  seeds  on  the  other,  only  in 
the  fact  that  spores  are  not  sex 
cells,  that  is,  do  not  need  to  imite 
with  other  cells   in   order  to  de- 


velop, ;is  is  tJK'  e.ise  uilli  most 
eggs    and    .seeds. 

Si'()VT,\Ni:ors  (Jknkhatiov.  Tlie 
(liltVrence  l)etween  "spontjuwous" 
in  this  phrase  and  in  the  phra.se 
"s])ontaneous  action,"  as  of  an 
animal,  shoidd  not  lie  missed.  In 
the  latter  comH-ction  the  wortl 
has  nearly  it"  not  (|iiitr  its  orig- 
inal meaning,  that  is,  of  one's 
own  accord,  or  initiativt*;  acting 
by  and  through  one's  self  alone. 
The  adjective  pronouns  men,  tun 
and  Hui'i  are  said  to  have  in-en 
used  always  with  fpojite  in  good 
Latin  ]>ro.se.  Strictly,  then,  if 
life  really  originated  from  some- 
thing which  was  not  living,  that 
is  by  a  "fortuitous"  concourse  or 
interaction  among  chemical  ele- 
ments of  different  sort,  xpnn- 
tnneoxt.s  would  not  be  the  proi>er 
term  to  descril)e  the  operati<Mi, 
simply  because  it  would  have  in- 
volved fundamentally  several 
selves,  even  if  the  different  ele- 
ments could  each  be  called  a 
.self.  It  would  not  have  been  an 
operation  itlentifiable  l)y  uni. 
your,  or  ///,«  or  its,  but  by  their. 
Plurality  rather  than  singularity 
of  action  would  be  the  essence 
of  the  conce])tion. 

St>:hkothopic.  Reacting  to  stinndi 
of  contact  with  solid  objects. 

Stoi.on'.  .\  prolongation  of  tin* 
body  of  some  plants  and  animals 
that  gives  rise  to  new  individuals 
by    btnlding. 

SiDoHii'AHors.    Sweat-}>roducing. 

.SvMinoTic.  Pertaining  to  a  state 
of  living  together  of  two  dissim- 
ilar organisms  fo  the  ;idv;intage 
of    botli. 

Synaptic.  In  cytology  ptrlaining 
to  siiuitjtfis,  tl>e  conjugation  of 
ehromosonu's  in  s«'x  cells  precetl- 
ing  the  reduction  divisions  con- 
nected with  the  maturation  of 
germ   cells. 

SvNCVTir.M.  As  usj'd  in  this  book, 
a   cytological    ti-rm    applied   U\   a 


388 


Glossary 


protoplasmic  mass  containing 
many  nuclei,  but  not  set  off  into 
distinct  cells.  The  entire  em- 
bryo is  of  this  character  in  some 
animals.  In  fact  a  few  embry- 
ologists  have  contended  that  dur- 
ing the  embryonal  stages  of 
most,  if  not  all  animals,  the  cells 
are  connected  by  protoplasmic 
strands  and  bridges,  making 
them  syncytia.  The  undoubted 
wide  prevalence  of  syncytial 
structure  among  animals  espe- 
cially, has  been  used  as  an  argu- 
ment against  the  cell-theory. 
Synthesis.  From  the  organismal 
standpoint  not  many  terms  used 
in  biology  are  more  important 
than  this.  The  etymological 
meaning,  placed  or  put  together, 
expresses  onh^  a  part  of  the  to- 
getherness of  an  organism;  the 
part,  namely,  which  pertains  to 
the  assimilative  activity  per- 
formed by  the  organism  on  its 
nutritive  substances.  This  proc- 
ess may  be  regarded  as  a  syn- 
thesizing one  in  nearly  the  lit- 
eral sense  (though  even  here  the 
process  is  more  one  of  self-ac- 
tivity and  less  one  of  external 
agency  than  seems  to  be  implied 
in  the  original  word).  But  it  is 
when  we  come  to  consider  the 
original  nature  and  power  of  the 
organism  by  virtue  of  which  it 
assimilates  food,  that  the  inad- 
equacy of  synthesis,  except  in  a 
much  modified  sense,  comes  to 
light,  for  the  organism's  ability 
to  assimilate,  that  is  to  put  or 
place  together,  its  nutritive  sub- 
stances is  wholly  dependent,  so 
far  as  we  have  evidence,  on  the 
fact  of  its  being  already  and 
originally  a  together  entity.  An 
organism  is  able  to  put  together, 
or  synthesize,  its  food  just  be- 
cause it  itself  is  together,  or  syn- 
thesized. A  synthesized  state  is 
a  prior  condition  to  synthesizing. 
To  be  an  organism  at  all  is  to  be 


synthesized. 

Systematic.  Pertaining  to  a  sys- 
tem; literally  a  standing  or  being 
together.  It  is  unfortunate  that 
"systematic"  has  come  to  be  re- 
stricted in  its  application  in  re- 
cent biology  to  the  formal  classi- 
fication of  plant  and  animal 
species.  As  a  matter  of  fact  a 
necessary  consequence  of  the 
unity  of  all  phenomena  of  the 
living  world  is  that  all  these 
phenomena  "stand"  in  some  nat- 
ural and  ascertainable  relation 
with  all  other  phenomena,  so 
that  all  biological  knowledge 
whatever  must  of  necessity  be 
systematic  if  it  really  corre- 
sponds to  nature. 

Taxonomy.  Mode  of  arrangement, 
the  branch  of  biology  which 
deals  with  the  classification  of 
the  species  of  plants  and  ani- 
mals. 

Test.  As  used  in  zoology  and  bot- 
any, an  external  covering  or 
tunic,  usually  nearly  lifeless, 
tough  and  resistant.  Its  office  is 
mostly  protective. 

Thymus.  A  gland  of  internal  se- 
cretion found  in  the  neck  region 
in  all  vertebrates,  and  connected 
originally  with  the  gill  system. 

Thyroid.  One  of  the  most  impor- 
tant glands  of  internal  secretion, 
located,  as  is  the  thymus,  in  the 
neck  region,  but  connected  em- 
bryonically  with  the  pharynx 
rather  than  with  the  gills  proper. 

Trochophore.  a  larval  stage  in 
the  lives  of  many  marine  worms 
and  molluscs,  characterized  by 
being  well  organized  for  swim- 
ming by  means  of  cilia  variously 
disposed  on  the  surface  of  the 
body. 

Trypsin.  One  of  the  chief  "active 
principles,"  or  enzymes  of  pan- 
creatic juice.  It  splits  proteids 
into  simpler  compounds.  It  is 
produced  by  some  plants  as  well 
as  many  other  animals  than  man 


Glossary 


S89 


and   vertebrates. 

Tunic.  In  botany  and  zoolo/ry, 
any  well  differentiated  membran- 
ous covering  of  an  organ  or  an 
entire  organism ;  mueb  tbe  same 
as  a  test. 

Tunicate.  Name  of  a  group  of 
marine  animals,  most  sharply 
characterized  by  the  cellulose- 
containing  tunic,  or  test,  which 
envelopes  the  body;  by  the  pe- 
culiar basket-like  respiratory  sys- 
tem; and  by  the  notochord  or 
precursor  of  the  vertebral  col- 
umn, possessed  by  all  the  species 
in  the  embryonal  life,  and  by  a 
few  during  the  whole  life,  fre- 
quently used  synonymously  with 
Ascidian,  which  see. 

Vagal.  Pertaining  to  the  vagus 
nerves,  one  of  the  tenth  pair  of 
cranial  nerves  in  all  true  verte- 
brates. 

Vaso-constrtctor.  Applied  to  the 
nerves  which  cause  contraction 
of  the  walls  of  blood  vessels. 

Vaso-dtlator.  Applied  to  nerves 
which  cause,  or  more  exactly, 
permit  a  widening  of  the  blood 


vessels  by  (liiniiiisiiing  the  tonus 
of  the  musclrs  of  tin-  vrssel 
walls.  Since  the  smaller  blood 
vessels  are  all  supplied  with  iKith 
constrictor  and  dihitor  nrrvrs 
the  constant  balancing  between 
these  nntngonistic  influences, 
l)oth  kinds  of  im])ulse  In-ing  in 
rcsjionse  to  the  general  needs  r>f 
the  organism,  this  sclK-me  illus- 
trates well  .'i  principle  of  equili- 
bration widely  operative  in  the 
animal  kingdoni. 

ViscKRAi..  A  term  used  in  zoology 
to  indicate  not  only  the  totality 
of  internal  organs,  but  also  the 
side  of  the  animal  on  which 
these  are  situated. 

Zygote.  A  body  formed  by  the 
conjugation  of  two  reproductive 
cells,  called  gametes.  Gametes 
and  zygotes  may  be  either  uni- 
cellular organisms,  or  the  repro- 
ductive cells  of  multicellular  or- 
ganisms. 

Zvm(k;kn.  The  en/.yme-producing 
substance  in  the  .secretory  cells 
of  glands  the  secretions  of  which 
contain  enzvmes. 


INDEX 


Alxlorlinlden,  K.,  i,  103  AiiKi'b.i,  (•()ni|)loxity   of,   i,   ?<*<);   to 

Al).sohitisiii,   in('t;i])livsicnl,    ii,    1.'>1  m.in,    2it\  ;    as    orpaoism    and    as 

Acacia,  ii,  f)})               *  cell,  297 

Accidental     products    of    chemical  Amours,   of   fishes,    ii,    2M 

change,  i,   111  Amphiljia,  i,  JOt 

Acorns,    storing   of,    by    woodpeck-  Ampliioxns,      on      isolated      ll|a^t<^- 

ers,   ii,   -21)9  meres     of,     i.     Jot;     on     u'ridfil 

Acromegaly,  ii,  134,  137  series  in,   ii,    10(1 

Action,    autocatalytic,    ii,    107  Analysis,    in     biological     reasoning. 

Action-system,   ii,   216  ii,  2(Hi;  and  al>straction,  y.W -,  arid 

Activity,      adaptive,      excessiveness  synthesis,  33.5;   remarks  (ui,   TM'f. 

of,  ii,  257  of  organic   substance,  311 

Activities,   instinctive,   tendency  to  Anatomist,   of   protozoa,    i,   2xti 

excessiveness  of,  ii,  3>f),  3()8  Ancestral,    i,    290 

Adaj)tation,   of   neural    activity,   ii,  Anger,    ii,    133,   317 

183  Animal,   odors  of,  i,  fti;   behavior. 

Adaptive  parallelism,  i,  335  ii,    30H;    human,    3S4;    kingdom. 

Ada])tiveness,     ii,     311;     of    subra-  3S1 

tional   j)sychic  activities,  350  Animals,  as  analytical   chmii^ts  of 

Adrenal   glands,   and   nervous   sys-  one   another,   i,   SS 

tern,  ii,   131  Annelids,   ii,    lf)3 

Adrenaline,     chemical     composition  Antagonisms,    ii,    175;    within    au- 

of,  ii,   133  tonomic  nervous  systt-m,  I7S;  co- 

Adrenin,   in   blood,   ii,   131;   effects  operative,  17S 

on   fatigued  muscles,  181  Anthropologist,   ii,   385 

Adsorption,  meaiiing  of,  ii,  344  Anthropology,  ii,  337 

Adventitious  buds,  i,  38  Anthropomorphism,  ii.  301 

Aggregation,   principle   of,   i,    183;  Anti-bodies,  i,  I0(i 

of  protozoa,  368;   and   synthesis,  Anticipatoriness,    ii,    2X2 

ii,  236                                    '  Antigens,    i,    100 

Air,  in  relation  to  consciousness,  ii,  Ants,    itehavior    of,     ii.     357,    37!J; 

290;  breath,  and  spirits,  338  larval,    spimuiig    of    co<-«xui    l»y. 

Albumen,  living  and  dead,  distinc-  311 

tion   between,  i,   78  Aphids.    i.   353 

Alchemy,  ii,  388  Ai)prn(licidaria,    ii.   3 

Alcvonaria,    ii,    97  Apperception,  rehitiori  to  tropisms, 

Aldrich,   T.    H.,   ii,   133  ii,     331;     definition     by     Wundt. 

Alga,    i,    41  333;    n-al    nature-   of.   3V3 

Allen,    B.    M.,    on    origin    of    sex-  Apphs.  odor  of.  i.  87 

cells,    i,    73;    on    removal   of   thy-  Argon,    ii.   311 

roid,  ii,   145                                     '  Aristotle,   i.   3;    ii.   J7S 

Alternation  of  generations,  i.  31(»  Arrhenius.  S..  i,   101 

American    biologists    and    organis-  Art,   creativrness   in.   ii,   J2.\;   crea- 

mal  theory,  i,  11  tivc   imi>ulsc  in,   337 

391 


392 


Index 


Artificial  parthenogenesis,  i,  345 

Ascidian,  bud  propagation  in,  i, 
50,  309;  egg  development  in,  ii, 
17;  tentacles  of,  98;  ganglion 
of,  187 

Assimilation,  ii,  205 

Association  in  psychology,  objec- 
tive and  subjective  sides  of,  ii, 
230 

Associationism,    ii,    228 

Associationists,  ii,  221 ;  and  ele- 
mentalists,   228 

Atomistic  theory,  ii,  151 

Atoms,  ii,  149,  160 

Attention   and  choice,   ii,  231 

Attitudes,  difference  between  sci- 
entific and  philosophic,  ii,  307; 
elementalist,    and    emotions,    321 

Attributes,  correlation  of,  i,  215; 
ii,  202;  physical  or  material,  and 
psychical  or  spiritual,  215;  struc- 
tural and  functional,  277;  eth- 
ical, 284;  observed  corporeal, 
289;  physical  and  chemical,  305; 
latent  of  oxygen,  341 ;  latent, 
343 

Autocatalytic  action,  factor  in 
growth,   ii,    107  . 

Automaticity,   ii,  221 

Autonomic  nervous  system,  ii,  128; 
vagal  or  cranial,  129;  sympa- 
thetic or  thoracico-lumbar,  130; 
sacral,  130;  antagonisms  within, 
178 

Avoiding  reactions,  ii,  252 

Axioms,  ii,  297 

Axones,  ii,   170 

Baboon,  i,  98 

Bacillus   biitschlii,  i,   262 

Bacteria,  i,  310;  membrane  and 
surface  structure,  257;  un- 
doubted organisms,  263;  classifi- 
cation of,  266 

Balance  of  organs,  i,  7 

Balanoglossus,  i,  223 

Balfour,  F.  M.,  i,  268 

Ballowitz,  E.,  ii,  2 

Barker,  L.  F.,  on  interrelation  be- 
tween internal  secretions  and 
nervous  system,  ii,  130,  138 

Basedow's  disease,  ii,  138 


Bateson,  W.,  i,  22 

Bayliss,  on  accidental  chemical 
products,  i,  112;  and  Starling, 
ii,  120;  on  meaning  of  autonomic 
nervous  system,  129 

Bearers,  i,  306;  of  heredity,  338 

Bees,  honey,  ii,  268 

Behavior,  complexity  of,  i,  289; 
animal,  ii,  208,  210,  227;  be- 
havior-knowledge, 277 

Behring  Sea,  ii,  211 

Benda,  C,  ii,  35 

Benecke,   W.,   i,   258 

Bernard,    Claude,    ii,    149 

Bichat,  i,  4 

Bio-chemical  substances,  phylog- 
eny  of,  i,  110 

Bio-chemistrv,  and  taxonomist,  i, 
94 

Biococcus,  i,  22;  319 

Biogen  conception,  i,  194 

Biogenesis,  theory  of,  i,  27;  versus 
spontaneous  generation,  316 

Bio-integration,  types  of,  ii,  94 

Biologist,   anthropological,   ii,   335 

Biology,  real,  i,  5;  goal  of,  ii, 
152;  elementalist,  and  associa- 
tionist  psychology,  228;  subdivi- 
sions of,  283 

Birds,  high  flight  of,  ii,  258;  song 
habits  of,  260;  mating  habits  of, 
263 

Blastomeres,  i,  203;  position  in  the 
whole,   206 

Blastula,  i,  203 

Blepharoplast,  i,  255,  329 

Blood,  and  bloods,  i,  91 ;  cor- 
puscles, white,  297;  adrenin  in, 
ii,   131 

Blue-jay,  storing  habits  of,  ii,  270 

Body,  i',  321;  ii,  150,  215;  relation 
to  mind,  216;  constitution  of, 
289;  vs.  corpse  or  cadaver,  322; 
and  soul,  323 

Born,   G.,   i,  207 

Botanical,  diagnosis,  i,  265 

Botany,  elementary  instruction  in, 
i,  236 

Boyle,   Robert,  ii,  288 

Brain,  not  coordinating  center,  ii, 
191;  normality  dependent  on, 
194;  as  element  of  organism,  216 


Index 


393 


Brandt,  Percy,  i,  30 

Bread,  "secret  powers"  of,  ii,  300 

"Breath     of     Life,"     ii,    303,    33(), 

338 
Brown,  A.   P.,  i,  95 
Briicke,  E.,  conception  of  the  cell, 

i,  129 
Bud,  adventitions  in   plants,  i,  38; 

propajration    in   com])ound   ascid- 

ian,   50,  309;   in   l)ryozoa,  53 
Burrows,  M.  T.,  on  tissue  cultures, 

i,    173;    on    organ    formation    in 

such  cultures,  176 
Butterflies,  larvae  of,  ii,  239 

Calkins,  G.   N.,  i,  240 

Canihiutn,  huds   from,  i,  39 

Canary    birds,    ii,    261 

Canidae,  i,  96 

Cannon,  W.  A.,  on  chromosomes 
and  Mendelism,  i,  356;  on 
heredity  in  plant  hairs,  ii,  55 

Cannon,  W.  B.,  on  autonomic  nerve 
action  and  adrenin,  ii,  129,  131, 
162,   178,   185,  319,  323 

Carbon,  ii,  338 

Carrel,  Alexis,  on  tissue  cultures, 
i,  168,  174;  isolated  tissues  and 
"morphological  plan"  of  organ- 
ism,  177 

Carrying,  characters  of  adult,  i, 
224;  hereditary  qualities,  second- 
ary and  acquired,  ii,  67 

Cassia,  ii,  99,   106 

Casteel,  D.  B.,  ii,  9 

Castle,  W.  E.,  definition  of  hered- 
ity, i,  315 

Caterpillar,  ii,  239 

Caudal  tube  of  spermatozoan,  ii, 
7 

Causal   factors,  ii,   103 

Cause,  in  heredity,  i,  313;  sufficient, 
ii,  147;  contributing,  153;  un- 
known, of  experience,  303 

Cell,  as  chemical  laboratory,  i,  82; 
physical  chemical  concej)tion  of, 
116;  as  an  organism,  Briicke, 
129;  as  key  to  ultimate  biologi- 
cal problems,  163,  181;  in  logical 
and  factual  aspects,  228;  as 
elementary  organism,  228;  ag- 
gregations,    295;     colonies,    295; 


evolution     of,     Minrhin,     K.      \  . 
307 

('«ll-meinbr.'me,  produced  by  pro- 
toplasm,   ii,    59 

C<'ll-nucleus,  and  protoplasm,  i, 
126 

Cell-state,   i,    295 

Cell-system,   i,   216 

Cell-theory,  what  it  is,  i,  150  e>l 
scq.;  inadequacy  of,  11  and  158; 
attempt  to  siiltordinalc  jirotista 
to,  280  et  seq.;  and  integration 
of  nervous  system,  ii,   16J) 

Cell-wall,    in    higher    plants,    ii,   51 

Ceils,  suliordiiiate  to  li\it)g  beings, 
i,  293;  used  bv  living  iK-ings, 
294;    isolated,    294 

C'ellular  centers,  i,  332 

Centers,  dynamic  in  cells,  i,  3:1:1; 
of  appercejition,   ii,   .'33 

Centrioles,   i,  :i33 

Centrosome,  i,  3:J0,  'Xi\ 

Cerebral   cortex,   ii,   21  (» 

Cerebrum,    ii,    130 

Ceremonies,  s<'lf-exhausting,  of 
mating,  ii,  264 

Chu'tognatha,    ii,    281 

('Jurtoptents,  i,   12 

Chain    reflexes,   ii,    197 

Chalh'nger  Exjiedition,  ii,  7 

Characters,  s))ecial  and  general 
in  heredity,  ii,  40 

Chemical  action  and  interaction,  i, 
215;  autocatal}  tic  in  organic 
growth,   ii,   105 

Chemical  basis  of  gemis  and  spe- 
cies,   i,    107 

Chemical,   criterion    of,   ii,   289 

Chemical  messengers,  i,  23;  ii,  119, 
121,   128,   170 

ClK'niico-functional  integration,  ii, 
94 

Chemico-naturalist  inijuiries,  I,  105, 
109 

Chemist,   and    naturalist,    i,    107 

Chemistry,  and  organism,  i,  75;  of 
organisms,  91;  comparative,  105; 
aiul  variation,  115;  in  solving 
jiroblems  of  heredity,  ii.  42; 
(Iclinition  of,  287;  i)liysical,  303, 
336;  periodic  law  of.  3?!);  atom- 
istic, 343 


394 


Index 


Chicken  pox,  i,  264 

Child,  C.  M.,  and  physiological  cor- 
relation, i,  17;  and  metabolic 
gradients,  ii,  108 

Chipmunk,  storing  habit  of,  ii,  271 

Chondriosome,  ii,  36;  as  material 
substratum  of  different  tissues, 
39 

Chromatin,  theory  of,  i,  314;  rela- 
tively undifferentiated,  318;  evi- 
dence of,  as  hereditary  sub- 
stance, 326;  physicial  basis  of 
heredity,  328;  supposed  omnipo- 
tence in  heredity,  ii,  14;  kinds 
of,  67 

Chromatinists,  i,  319 

Chromosomal  elementalism,  i,  320 

Chromosomal  hypothesis  of  hered- 
ity, evidence  for,  i,  324,  326  et 
seq. 

Chromosome  dogma,  ii,  59 

Chromosomes,  i,  21,  306,  324;  as 
immediate  ancestors,  319;  in 
fertilization,  342;  accessory,  347; 
X  and  Y,  350;  seat  of  inheri- 
tance material,  ii,  22;  in  rela- 
tion to  heredity,  66;  initiators  in 
heredity,   83 

Chun,   C,  i,  301 

Cilia,  i,  330 

Ciliary    tuft    of   spiriUum,    i,    259 

Classification,  i,  99,  296;  of 
physical  facts,  ii,  217;  synoptic, 
276 

Chemical  evidence  of  adrenal- 
nervous  connection,  ii,  133 

Cohn,  F.,  on  classification  of  bac- 
teria,   266 

Collins,  H.  H.,  ii,  258 

Common-paths  in  nerve  physiol- 
ogy, ii,   171 

Common-sense,   i,   32 

Comparison,  i,  99;  of  shells  of 
rhizopod  and  nautilus,  237; 
sacrifice  of,  in  experimental 
method,  ii,  279. 

Competition,    ii,    175 

Condition,  molecular  appeals  to, 
i,  276 

Conjugation,   i,   269 

Conklin,  E.  G.,  on  eg^  as  stage  in 
life     of     organism,     i,     193;     on 


development  of  Ascidian  e^^,  ii, 
17;  on  hereditary  characters  de- 
termined by  cytoplasm,  and  by 
chromatin,    42,    et    seq. 

Coordination,  neural,  not  a  "cen- 
tral"  process,   ii,   192 

Corycella,   i,   270 

Consciousness,  ii,  161;  contents  of, 
233;  organismal  theory  of,  282; 
and  chemical  action,  290;  theory 
of,  and  theory  of  knowledge, 
296;  an  attribute  of  the  organ- 
ism as  a  whole,  309;  and 
physico-chemical  conception  of 
organism,  324;  and  pro-con- 
sciousness,  350 

Contents  of  consciousness,  ii,  225, 
233 

Courtship    of    animals,   262 

Cowdry,   E.   V.,  i,  437 

Crampton,  H.   E.,  ii,  26 

Crane,  sand-hill,   ii,  258 

Crephlula,  ii,  20 

Cretin,   ii,    116 

Crickets,   chirping   of,   ii,   261 

Cnthidia,  i',   334 

Ctenophore,    i,    201 

Cushing,  H.,  ii,  113,  124 

Cuvier,  i,  5 

Cycads,    ii,    58 

Cytoplasm,  and  Karyoplasm,  i, 
135;  Kinds  of,  ii,  67;  funda- 
mental and  primitive  as  heredi- 
tary   substance,    68 

Cytoplasmic  activity,  in  spicule 
production,    ii,    52 

Cytoplasmists,   i,   319 

Cytostome,   i,   248 

Dances,  of  lapwing,  ii,  262 

Darwin,  Chas.,  as  naturalist,  i,  75; 
as  example  of  creativeness  in 
science,  ii,  225;  on  comb  of  hive 
bee,  168 

Davidson,  H,  C„  on  plant  as  sym- 
biotic colony,  i,  35;  on  "planto- 
gens,"  36 

De  Bary,  and  cell  theory,  i,  162 

Definition,   i,   296 

Dendrites,  ii,   170 

Descartes,  Rene,  ii,  298 

Descent,    i,    315 


Index 


'Ml 


I^escription,   i,  J)9 

Deterniinaiits,  Weisinannian,  i,  21, 
^2o\  changed  to  determiners,  3iH 

"Determined,"  different  meanmgs 
of,    ii,    49 

Determiner,  faseination  of  for 
some  minds,  i,  306;  meaning  in 
true  objeetive  sense,  ii,  Ki;  and 
dialeetics,  ?(>;  theory  of,  con- 
trary to  eheinieal  principles, 
79;  \\'ilson's  proposal  to  drop, 
82 

Develo])ment,  in  ])roto/.oa,  i,  2()7; 
cause  of,   ii,   158 

Developmental    mechanics,    i,    18 

Dewey,  John,  ii,  298;  on  "Self" 
and   environment,  305 

Diagnosis,   medical,   i,   ^{iC) 

Dialectics,  and  determiner  hy- 
jiothesis,   ii,   7G 

Diatoms,  i,  310 

Difference,  chemical,  between  or- 
ganisms, i,  83;  between  germ- 
cells,  importance  of,  214;  re- 
semblances and,  317;  in  func- 
tion  and   l)ehavior,  ii,   27() 

Differential    factor,    ii,   82 

Differentiation  and  integration,  ii, 
1()8 

DinoHagellates,    i,    310 

Directing  activity,  of  develoj)- 
mental  process,  i,  70 

Dis]>lay,    mutual,    ii,   264 

Distribution,    vertical,    ii,    281 

Division,  of  labor,  i,  205;  deter- 
mined by  growth,  220;  })hysio- 
logical,  ii,  24 

Dobell,  C.  C,  on  nuclei  in  bac- 
teria, i,  262;  on  Ehrenberg's 
conception  of  protozoa,  284;  on 
protozoa    as    non-cellular,    290 

Dog,  as  causal  explanation,  ii, 
203,  204 

Donaldson,  H.   H.,  ii,  169 

Doncaster,    I..,    i,   352 

Dormitive    j)rinciple,    ii,    204 

Dramatist,   ii,   317 

Driesch,  Hans,  on  cell  theory,  i, 
153;  totipotence  theory  (»f,  202, 
et    seq. 

Ductless  glands,  ii,  114 

Dujardin,    Felix,   interj)retation    of 


protozoa,     i,     2H();     .md     plasunc 

(JcnH-ntalisni,    3Jo 
Duodenal     mucous     nu-nibrane,     ii, 

119 
Dynamic   (•enter,   of   cell,    j,   :VXi 

Earthworm,    ii,    191 

Hihinns.   i,  202 

Ideology,    ii,    212,    27!) 

I'.cononiy,  |)liysiol(>gic;il.  ii,   .»(i| 

]\ctoderm,  i,  46 

Kffect,    i,   313 

l">gg,  of  chick,  studied  clH-uiieallv, 
i,  79;  of  frog,  199;  tlo.iting,  213; 
h<'re(lit;iry  attributes  of,  JH;  ii.s 
stage  in  development  of  iiuli- 
vidual,   ii,   24 

I'Jirenberg,  C\  (1.,  interpretation 
of   j)r()tozoa,   i,   280,   and   ii,  «»6 

l^lenuMits,  photosensitive,  ii,  IS«>; 
physical  and  chemical,  235; 
psychical,  2'.].');  cheniieal.  cri- 
terion of,  286 

l\lementalism,  i,  2;  narrowing  in- 
fluence of,  230;  cellular.  2S(i; 
and    internal    secretion,   ii,    IH 

Klementalist,  conce})tion,  i,  2so; 
sjieculation,  319;  and  organismal 
standpoints,  ii,  11-S;  theory,  and 
neglect  of  fact,  157;  anarchistic, 
1(J0;  attemjjt  to  interj)ret  trop- 
istic  aiul  segmental  tln'ories, 
19S;  l)iology  and  associ.itionist 
])sychology,    22S 

Klementalistic  interpretati«»n,  ii, 
23 

KlenuMitary    organism,    i,    227 

1 '.nilu'yogeny,    i,    277 

Kmbryology,  nu-thods  of,  i,  221; 
of  protozoa,  268;  and  genetics, 
311,   324 

Embryo,  i,   201^,  272 

Emotions,  ajid  |)hysical  organir.a- 
tion,  ii,  21();  tuitural  history  de- 
scription of,  '.\\H,  322;  elenien- 
talist    descripliiui   of,  :i2I 

Emotional,  attitucU-.  i.  322;  (ilye<>- 
suria,  ii,  13.*;  psychic  life  of 
animals,    i:{:{ 

Em|)e(locUs,   i,  3,   40 

I''iuioderni,    i.    Mi 

I'lidocrin**  glands,  ii,   I  I  I.   KM) 


396 


Index 


Endoplasm,    i,    277 

Energy,  formative  and  regenera- 
tive, i,  11;  of  modern  physics, 
76;  and  matter,  141;  conception 
of,  320;  of  contraction,  ii,  63; 
and  substance,  337;  and  power, 
force,  work,  342;  surface,  344 

Entelechy,  ii,  149 

Environmental  influence,  ii,  244 

Enzymes,  digestive,  comparative 
chemistry  of,  i,  104;  facilitate 
transformation,  ii,  81 

Epicurus,  i,  3 

Epithelium,  and  internal  secre- 
tions, ii,  135 

Eppinger,   H.,   ii,   133 

Equilibrium,  general  notion  of,  i, 
17;  physical  chemistry  concep- 
tion  of,   216 

Essences,  ii,  288 

Esterly,  C.   O.,  ii,  208 

Evidence,  direct  and  indirect,  of 
mechanism  of  heredity,  i,  325; 
favorable  to  chromatin  as  hered- 
itary substance,  326;  visible  and 
invisible,  ii,  5Q\  picking  out  of, 
ii,   263 

Evolution,   i,   291,   ii,   241 

Evolutions,  four  simultaneous,  1, 
321 

Eudendrium,   i,   67 

Eugenics,  i,  305;  and  fatalism,  ii, 
89 

Excessiveness,  of  instinctive  activ- 
ity, ii,  259;  of  sex  impulse,  265 

Excitability,  threshold  of,  ii,  165; 
selective,  165 

Excitor,   ii,    147 

Experience,  ii,  305;  subjective  and 
objective,  285;  causes  of,  un- 
known, 303 

Experimentation,  limitations  of,  ii, 
212;  laboratory,  279 

Explanation,  spurious  type  of,  ii, 
81;  causal,  146,  240;  of  trop- 
isms,  190 

Expression  of  emotion,   ii,  264 

Factorial  hj^othesis  in  heredity,  i, 

21 
Factors,   in   heredity,   i,   42,   306 
Facts,  neglect  of,  by  elementalists, 


ii,  157;  matters  of,  299 

Falta,    W.,    ii,    134 

Fatalism,  ii,  89 

Fatigue,  and  sugar  in  blood,  ii, 
132;  and  blood  pressure,  181 

Fear,  and  adrenin  in  blood,  ii, 
132;  with  physical  and  psvchical 
life,  183 

Feeling,  intellect,  will,  ii,  217 

Felidae,  i,  96 

Ferns,  ii,  99 

Fertilization,  hybrid,  i,  344 

Fielde,  A.,  on  odors  in  ants,  i,  89 

Finalism,   ii,    152 

Finch,  house,  ii,  260 

Fishes,  sperm  of  diflferent  species, 
i,  102;  floating  eggs  and  young 
of,  213;  mating  habits  of,  ii,  265 

Flagella,  of  bacteria,  259;  rela- 
tion to  nucleus,  328,  330;  struc- 
ture  and  origin  of,  370 

Flat-worms,  ii,  109 

Flavors,  of  animals  and  plants,  i, 
84 

Fluid,  as   phase   of  system,  i,  216 

Foetus,  i,  272 

Forbush,    E.    H.,    ii,    270 

Forces,   antagonistic,  ii,   127;   con- 
stitutively       antagonistic,       134;^ 
abuse  of  the  term,  298 

Fore-foot,  horse's,  and  man's  hand,] 
ii,  224 

"Formative  stuffs,"  i,  16;  and  in- 
ternal secretions,  ii,  142;  oi 
Sachs,   147 

Form--determination,  cytoplasmic,! 
ii,  14 

Forsyth,  Ruth,  ii,  44 

Fossil   wood,  ii,  58 

Foster,  Sir  Michael,   ii,  346 

Fragmentation,  and  analysis,  ii, 
236 

Freud,  S.,  ii,  350 

Friedlander,   B.,  ii,   102 

Frog,  eggs  of,  i,  199;  pigment] 
cells  of,  339;  sex  determination! 
in,  ii,  76;  larva  and  thyroid  of,| 
143;    croaking   of,   261 

Fruit  flies,  supposed  connection! 
between  mutations  and  chromo-1 
somes  in,  i,  354 

Fundamental,  criterion  of,  ii,  201 


Index 


397 


Fungus,   i,  41 

Fur-seal,     (Irvi'lopmcnt     of    sperm 

of,   ii,   •!•;   inigratiuus   of,   ii,   2\()\ 

mating   lialiits   of,   2()7 
Fusibility    of    tissues,    as    test    of 

relationshij),    i,    WW 

Ciall-stones,    ii,    1(J4 

danietes,  i,  i?()9 

(langlia,  supra-frsopliageal,  ii, 
lf)l;  as   relay  stations,  191 

Ciastrula,    i,    2i)\\ 

elates,  R.   H.,  i,  W^W 

(lenniuile,  as  method  of  asexual 
proj)agation,  i,  309;  as  speeula- 
tive  entities  (see  term  in  glos- 
sary) 

Gene,  a  hypothetical  entity  in 
modern  genetics,  1,  20;  static 
nature  of  the  conception,  \2; 
historic   antecedent    of,   ii,   84 

Geneticists,  modern  school  of,  i, 
324 

Genetics,  tendency  to  ignore  em- 
bryology, i,  311;  inattention  to 
develo]Mnental  facts,  ii,  14; 
thinking  on,  12 

Germ,  nature  of,  i,  2i23;  imaginary 
structureless,   268 

German  Deep-Sea  FiXpedition,  i, 
278 

Germ-cells,  in  hydroids,  i,  60; 
promorphologv  of,  211;  "throw 
of"  the  soma,  319;  subject  to 
metabolism,   ii,   74 

Germ-disc,   i,   208 

Germinal,  continuity,  i,  310;  ma- 
terial, 209;  localization,  ii,   16 

Germ-layers,  i,  45;  subservient  to 
organism,  48;  and  the  germ- 
plasm   theory  of  Weismami,  58 

Germ-])lasm  theory,  type  effect  of, 
on  observation,  i,  W>-^  implica- 
tions of,  318;  extreme  form  of 
contrary  to  inductive  science,  ii, 
76 

Germ-regions,  organ  forming,  i, 
209 

Giardia,   i,   254 

Gibbs,   J.    Willard,    ii,   341 

"Gifts,"   natural,   ii,   235 

Ginkgo,  i,  330 


Gladness,  fits  r)f,  jj,  263 

Gl.uuls,  ductless,  etxIcM-rine,  ii,   lit 

(ilycogen,   i,   215 

Goal   of  biology,   ii,   152 

Goette,  Alcxarubr.  rr\i«w  and 
criticism  by  W'cismnnn  on  m-x- 
cells  in  hydroids,  i,  62,  et  tin.: 
organismal  trend  of  residts  bv. 
68 

Goitre,   ii,    115 

Gradients,  (brect  and  in\crsr,  ii, 
106;   axial    ni«-tab(»li<-,    107 

Grafts,    i,    \V.\ 

Graiuil<s,    epithrbal.    ii,     13.) 

GrassliopjuT,   i,  :i57 

Graves'  disease,  ii,   133 

CJrebe,    great    crested,    ii,    263 

Grenricll,   .Joseph,    i,   85 

Groos,   Karl,  ii,  27.3 

Growth,  determines  division,  i, 
220;  integration,  ii,  93,  94;  cycle 
of,  105;  of  an  organism,  105; 
explanation    of,    107 

(Judernatsch,  J.    F.,   ii,    1 13 

Habits,  mating,  of  l)irds,  ii,  2»)3; 
of  viviparous  fishes,  2i\5\  st<ir- 
ing,  of  honey  bee,  268,  of  wood- 
pecker, 2()9,  of  mammals,  271; 
in    nature,    278 

Haeckel,  1'.,  on  rndiolnria,  i,  2'WJ\ 
moneron    theory,    256,   320 

liaecker,  ^^,  on  radiolaria,  i,  2*36; 
on    ontogeny    of    radiolaria,    278 

Hairs    of   higher    plants,    ii,   55 

Half-embrvo,    i,    199.    201 

Hallez,    P.,'  i,    218 

Hardens,    A.,    i,    lOt 

Hargitt.   C\   \V.,   i,   67 

Harnier.   S.,    i,   54 

Harmonic,  etpiijxttential  system, 
i,    205;    e(|uilil)riun),    ii,    118 

Harmony,    i,    3;    in    iM-alth,    ii,    l-'7 

Harrison,  H.  G.,  on  tissue  cul- 
tun-s,   i,    16S,   et   teij. 

Heidenhain,    H.,   i,  33? 

Heirike.    Fr.,   i,   213 

Heliotropic,   ii,    240 

Helium,   ii,  3t2 

HemoLdobin,    i,   95  * 

Henderson.    I..    J.,    ii.   339 

Ilerbnrt,    ns    extremi'^t    in    associ»i- 


398 


Index 


tionist  psvchologv,  ii,  229 

Herbst,   C.,'i,  344  ' 

"Hereditary  substance,"  contrary 
to    facts    in    hydromediisae,   i,   68 

Heredity,  and  elementalistic  phil- 
osophy, i,  20;  nature  of,  305; 
stronghold  of  biological  elemen- 
talism,  305;  effort  to  restrict  to 
sexual  propagation,  308;  defined 
by  E.  G.  Conklin,  308;  com- 
plex of  causes,  313;  definition, 
314;  chromatin  in,  320,  321;  and 
sex,  348;  cytological  basis  of, 
349;  spermatozoon's  tail  mani- 
festation of,  ii,  3;  mitochondrial 
theory  of,  33;  and  sponge  spic- 
ules, 53;  summary  of  informa- 
tion on  physical  basis  of,  64; 
narrowing    definition    of,    85 

Herlitzka,   Amedeo,   i,   205 

Hertwig,  O.,  theory  of  Biogene- 
sis of,  i,  27;  experiments  on 
half  embryos,  200;  on  centro- 
some,  332 

Hertwig,  R.,  on  cell-theory  as  ap- 
plied to  protozoa,  i,  288;  on 
experimental  determination  of 
sex,   ii,    76 

Higher  Usefulness  of  Science,  ii, 
337 

His,  Wilhelm,   1,  208 

Histogenesis,  and  the  mechanism 
of  heredity,  i,  325,  and  ii,  32; 
and  species  characters  in  adults, 
43 

Hobbes,  Thomas,  and  sensational- 
ism in  philosophy,  ii,  219 

Holmes,  S.  J.,  an  organism  as 
symbiotic  community,  i,  183;  on 
brainless  frogs,  ii,  195;  on  ac- 
tivities of  Amphithoe,  248;  on 
behavior  of  ants,  257 

Hooker,  D.,  i,  339 

Hopkins,  F.  G.,  on  the  cell  as  a 
chemical  laboratory,  i,  82;  on 
the  physical  chemistry  of  the 
cell,  114  et  seq.;  on  the  cell 
constituents  essential  to  the  cell 
as  a  system  of  phases,  192;  ver- 
sus' particular  types  of  mole- 
cules as  an  explanation  of  life, 
194 


Honey-bee,  eggs  and  chromosomes 
of,  i,  352;  storing  habits  of,  ii, 
268 

Hormones,  importance  of,  to  or- 
ganismal  conception,  i,  23;  na- 
ture of  action  of,  ii,  121 ;  rela- 
tion to  nerve  action,  128;  sup- 
posed identification  with  "form- 
ative stuffs,"  142;  integrative 
office  of,  compared  with  that  of 
nervous  svstem,  162 

Hudson,  W!  H.,  ii,  261 

Human  being  in  one-celled  stage, 
i,   217 

Hume,  David,  ii,  298 

Hunter   ciliates,  i,  235 

Huntsman,  A.  G.,  ii,  44 

Huxley,  Julian   S.,  ii,  263 

Huxley,  T.  H.,  on  the  physical 
basis  of  life,  i,  121 ;  on  the 
cell-theory,  288,  296;  on  innate 
ideas  as  conceived  bv  Descartes, 
ii,  298;   against  materialism,  302 

Hydroids,  germ-cells  in,  60;  sepa- 
rated blastomeres  in  eggs  of, 
204;  graded  growth  series  in,  ii, 
97 

Hypophysis,  ii,  124;  alliance  with 
thyroid    and    adrenals,    127 

Hypopituitarism,   ii,    113 

Hypothesis,  ii,  282;  of  conscious- 
ness,   286;     "working,"    291 

Idea,  central  of  this  book,  i,  24 
Ideas,     atomistic     and     association 
of,    ii,    229;    "relations    of,"    in 
Hume's  system,  299;  innate,  301 
"Identical  stuffs,"  i,  123 
Impertinence,   scientific,   ii,   247 
Individual,  man,  i,  31 ;  tree,  31 ;  ex- 
altation  of,   196;   normal,  ii,  205 
Individuality,   in   the   living  world, 
i,  30;   Huxleyan,  43;  of  chromo- 
somes, 85;   of  organism,  ii.  111 
Inheritance,  i,  312;  nuclear  theory, 
and    cytoplasmic    localization,    ii, 
2-2 -^    of   acquired   characters,   24; 
material — imitator     rather     than 
determiner,   QG\  probability   that 
substance  becomes   such   in   each 
ontogeny,  73 
Inhibition  of  reflexes,  ii,  176 


Index 


399 


Initiativi',  mental,   ii,  2\\ 

Initiator  hypothesis  in  hereditx, 
advantages    of,   ii,   8;i 

Inner  mass,  and  outer  la\cr.  r»f 
l)0(ly,    i,   :i01 

Insane-h'ke  eonthict  Irom  al)sencc 
of    I) rain,    ii,    \\)\ 

Inseet,  e^'^s  of,  i,  '220;  sperm  of, 
ii,   J) 

Instinct,  involves  animal  as  a 
whole,  ii,  1S8;  inijiortanee  of  as 
zoological  term;  247;  variability 
of,  251;  and  intelligence,  256; 
food-gathering,  2()8;  problem  of, 
284;  "instinct  actions,"  "instinct 
feelings,"  284;  and  physical  or- 
ganization,   310 

Integrratcdness,  of  Oinrdia,  i,  255; 
and   equilibrium,   ii,   198 

Integration,  growth,  ii,  93;  func- 
tional, 113;  distinction  between 
develoj)mental  and  functional, 
161;  cellular,  in  reflex  arc,  l(i.3; 
and  differentiation,  1()8;  ps\- 
chic,   214 

Integrity,    organismal,    i,    26 

Intellect,  feeling,  will,  ii,  217;  and 
instinct,    236 

Interaction,    chemical,    i,    215 

Intercellular  substance,  ii,  170 

Interde])endence,  metabolic,  ii,  104 

Internal  secretions,  importance  of, 
to  organismal  concejition,  i,  23; 
conception  of,  ii,  113;  interrela- 
tions of,  124;  and  epithelium, 
135;  and  "formative  stuffs,"   142 

Internal  secretory  systems,  ii,  128; 
and   nervous  systems,   128 

Investigation  of  distribution  of 
sex-cells,  i,  73;  statistical  of 
animal  behavior,  ii,  280 

Iron,    ii,   329,   337 

Ii-radiation,    ii,    17t 

Isolated    cells,   i,    167 


James,  William,  on  human  energy, 
ii,  132;  on  associationist  psy- 
chologv,  229;  on  consciousness 
of  self,  309 

Jelly-fish,  i,  235 

Jennings,  H.  S.,  on  ultra-.Mendel- 
ism,    i,    42;     on     com^jlcxity     of 


protozoan  behavior,  2H9;  on  trial 

and   error,   etc.,   ii,  252 
Johnson.    II.    I'.,    i,    272 
Jolly,    .1.,    i,    I7(» 
Jordan,    FI.    I''.,    ii,    60 

KanI,  Iv,  self-conscious  uiiitv  of 
apperc<ption,  and  t  ranseend«'H- 
talism,   ii,   233 

Karyoplasm,   i,    135 

Kelp,    ii,    105 

Key,  cell  as,  to  liiojogjeal  jilie- 
nomena,   i,   229 

King,  Miss    II.    I).,   ii,   76 

Knowledge,  analytic  and  synthet- 
ic, i,  23;  -getting,  ii,  213;  ana- 
Ivtic,  213;  theorv  of,  296;  nature 
of,    297 

Kofoid,  ('.  A.,  i,  241;  on  soil  amoe- 
ba, 32S 

Kofoid  and  Christiansen,  on  neu- 
rometer  system  in  flagellates,  i, 
J51 

Kolle   .ind    \\  assermaim,  i,   258 

Laboratory,   as    an    agency,    ii,    212 
I.amarek,   i,   5,   75 
I.ancelet,   ii,  95 
Langley,  J.  M.,   ii,   129 
Lapwing,  dance  of,  ii,  262 
Lar\;e,    grafting    together,    i,    207; 

specific    characters     in,    214;    ns 

"carrier"     of     adult     chnractcrs, 

224 
Law,   of  elements,    ii,    KiO 
Lead,    ii,   341 
Leaves,   compound,    ii.   99 
Leeches,    ii,    311 
I.eibnitzian    theory,   ii,    150 
Lemon    trees,    i,   38 
Le\\  is   and    Lewis,   ii,    10 
Life,     ii,    28(j;     veg«'tal,    emotioiud, 

rational     and      intellectual.     16.'; 

tripod    of,    18.';    subp'eti\e,    .N I 
Like   |)roduces   like,   i,  315 
Likenesses,    functional,    ii,    276 
Lilli«',    F.    IL,   as   ]>re-organismMlist. 

i,    11;    on    "j)roj)erti<'s    of   whole" 

in  the  embrvo,  12  and   193 
Lillie.    K.   S.,  'i.  310 
Linudus    nniseb',    ii,    (50 
Lineus    larlt  im,    i,    189 


400 


Index 


Living  beings,  in  nature,  ii,  211 
Living  substance,  i,  115 
Living  units,  hj'pothetical,  i,  19 
Localization,  by  protoplasmic  flow- 
ing,  ii,   19 
Locust,  i,  218 
Locy,    Wm.    A.,    i,    280 
Loeb,  Jacques,  i,  23;  on  relative  in- 
fluence of  nu  cleus  and  protoplasm 
on  heredity,  ii,  41;  on  identifica- 
tion   of   internal    secretions    with 
"formative      stuffs,"      141;      on 
"ultimate   aim"   of  biologj%   151 ; 
and  "organism  as  a  whole,"  185; 
neglect    by,    of    work    of    Sher- 
rington   and    Cannon,    185;    on 
understanding    of    natural    phe- 
nomena,    207;     organismal    ten- 
dency    of    tropism     theory     of, 
240 
Logic,  pure,  i,  22 
Loomis,  L.  M.,  i,  85 
Love,  emotions  of,  ii,  265 
Luciani,  L.,  ii,  115 
Lucretius,    i,    3 

Machines,   living,  ii,   252 

Macrocystis   pyrifera,   ii,   105 

Mammals,  storing  habits  of,  ii,  271 

Manly,  J.  M.,  on  "exuberant  vital- 
ity" of  Shakespeare,  ii,  223 

Marceau,  F.,  ii,  61 

Marshall,  F.  H.  A.,  ii,  79 

Materialism,  author's  attitude  to- 
ward, ii,  207;  Huxley  against, 
302 

Mathematics,  ii,  297 

Mating  habits,  of  birds,  ii,  263; 
of  fishes,  265 

Matter,  and  energy,  in  modern 
physics,  i,  76,  141;  and  force, 
196;  composition  of,  288,  341; 
generality  of  the  term,  304 

Mass  action,  ii,  344 

McClung,  C.  E.,  i,  347 

Mcllvane,  Charles,  i,  87 

McMurrich,  J.,  Playfair,  on  germ 
layers,  i,  47;  character  of  cell 
division  in  embryo,  219 

Meadow  lark,  western,  song  habit 
of,   ii,  259 

Mechanism  of  heredity,  i,  315, 
322;  organic  vs.  inorganic,  ii,  252 


Medussetta,  i,  236 
Meirowsky,    E.,    i,    339 
Melanin,   i,   339;    formed   in   cyto- 
plasm, 341 
Membrane,   and   surface   structure 
of  bacteria,  i,  257;  synaptic,  be- 
tween cells  of  reflex  arc,  ii,  167 
Mendel,  Gregor,  i,  305 
Mendelian    inheritance    and    chro- 
mosomes, i,  356 
Mendelism  as  a  creed,  i,  324 
Mental,  sense,  i,  3;  initiative  and 

restlessness,  ii,  243 
Meristic  parts,  ii,  95;  meristic  phe- 
nomena   in    plants    and    in    ani- 
mals, 103 
Merotomy,  i,  276 

Mesenchyme  cells,  dormant  in  tad- 
poles,    ii,     147;     as     inheritance 
material,  155 
Messengers,    chemical,    secretin    as 

example  of,  ii,  119,  121 
Mesoderm,  i,  46 

Metabolic    processes,    and    the    or- 
ganism's     supremacy     over     its 
cells,  i,  294;  interdependence  of, 
ii,  104 
Metabolism,  i,  215;  germ  cells  sub- 
ject   to,    ii,    74;    katabolic    and 
anabolic,    346 
Metals,  "base"  and  "noble,"  ii,  288 
Metameres,  ii,  95 
Metamorphosis,  ii,  145 
Metaphysician,   ii,   285 
Metaphysics,     chromatin     and,     i, 
321;  juvenile,  ii,  141;  as  an  epi- 
thet, 201 
Metaplasv,   of   diiferentiated   cells, 

i,   186  ■ 
Metazoa,  i,  268 
Metcalf,   M.    M.,   i,  289 
Method,  experimental,  ii,  278;  sta- 
tistical, employed  at  Scripps  In- 
stitution,   280;    natural    history, 
in    study    of    self,    282;    impor- 
tance  of,   282 
Meves,  F.,  ii,  35 
Meyer,  Arthur,  i,  258 
Mice,   summersaults   of,   ii,   258 
Michael,  E.  L.,  ii,  281 
Microbes,   specificity   of,   i,   265 
Miescher  and  Kossel,  i,  79,  102 
Migration,   of  sex-cells,  i,   61;   ex- 


Index 


401 


tent  of,  hv  birds,  ii,  QH^ 

Milk,  i,  lot" 

Mincliiii,  V,.  A.,  Bioooccus  theory 
of,  i,  2;  on  evolution  of  eell, 
307;   as  chrornatinist,  318 

Mind,  relation  to  liodv,  ii,  21.>, 
2U);    "mind    stuff,"   'M9> 

Miracles,  i,  322 

Mitochondria,  in  ontopeny  of  in- 
sect sperm,  ii,  9;  not  trans- 
formed   into    neuro-fibrils,   37 

Mitochondrial  theory  of  heredity, 
ii,  33 

Moeser,  W.,  ii,  107 

Molecular  condition,  appeals  to,  i, 
276,  282 

Molecules,  ii,  149 

Mollusca,  i,  221 

Moore,  V.  A.,  i,  259 

Monogamy,    in    viviparous   fishes, 
ii,   265  * 

Moneron  theory  of  Haeckel,  i,  256, 
320 

Morgan,  C.  T.loyd,  ii,  328 

Morgan,  T.  H.,  and  regeneration, 
i,  180;  on  half-embryos  of  frog, 
200;  on  eggs  of  phylhixerans, 
353;  on  mutations  and  heredity 
in   fruit  flies,  354 

Morphallaxis,   i,    180 

Morphological,  plan  of  the  organ- 
ism, i,  177,  195;  entity,  funda- 
mental to  tropism  conception,  ii, 
190 

Mortensen,  T.  H.,  i,  216 

Mosaic,  theory  of  development,  i, 
12,  198,  205;  "mosaic  picture," 
organism   as,  208 

Mosquitoes,  i,  213 

Mosses,  leaves  of,  ii,  57 

Mushrooms,  ]ioisonous  species  of, 
i,    87 

Musk-deer,  i,  86 

Mustard  plant,  weight  of  seeds  of, 
ii,   102 

Mutation,  connections  of,  with 
chromosomes,  i,  353 

Myomeres,   ii,   95,    106 

Mystification  of  protoplasm,  i,  121 

Natural  history,  special  ability  of, 
i,   113;   methods  of  studying  the 


self,  ii,  282;   importance  <.f,    ?<i:i 

Natural  selection,  and  JM-lectrd 
evidence,  ii,  263;  slight  rri:«r(l 
for  quantity  by,  259 

N'aluralism,    ii,    112 

Naturalist,  and  rvnbition  tlir<»rv, 
i,  76,  286;  ii.  2J1;  zoological, 
227,   278 

Nattire,  of  things,  i,  34;  not  sim- 
ple,   236;    of    knowledge,    ii.    1.',.' 

y<infiliin.  comparison  of,  with 
rhizopod,    i,   237 

Needs,    nutritional,    ii,    268 

Neglect  nothing,  naturalist's  mot- 
to, i,  91;   ii,  216,  245,   283 

Nemertean,  regeneration  of,  i. 
189;  capture  of  jirey   l)y.   ii,   P.V) 

Neoformation,    i.   276 

Kcrri.'i.  ii,  192 

Neresheimer,  E.  R.,  i,  245. 

Nerve,  centers,  ii,  187;  phvsiologv 
of,  320 

Nervous  .system,  in  proto/.oa,  i, 
243;  autonomic,  ii,  128;  and  in- 
ternal secretions,  128;  integra- 
tive action  of,  162  rt  sfff.: 
I.oeb's  important  conception  of. 
187 

Neural   integration,   ii,  9t,   161 

Neurones,  ii,  216 

N(Miroj)lianes,  i,  245 

Neiiro-motor  apparatus,  i,  243; 
system,  255 

Nickel,   ii,  327,   329 

Night-hawk,   ii,  258 

Nomenclature  of  germ-j)lasm  tlie- 
ory,  ii,  87 

Non-cellular,  jirotozoa  interpret«il 
as,  i,  290 

Nucleus,    i,    207,    'V.U  :    irj    bacteria, 

261;  part   in   pigm<'nt    formation, 

340;   in   oxidi/ing  action.  :U1  ;   of 

"utmost    theoretical   importance," 

ii,   22;   control   of  cytoplasm   by, 
1") 

Nusbaum,  J.,  on  ti'S'^ue  trans- 
formation,   i.    10(1 

Nusbaum  and  Oxner,  on  regrnern- 
tion   of   nemertean,   i.   189 

( )l)i<'cti\e,  side  of  psychical  ns5o- 
ciation.  ii.  2:^0;  and  "outer."  29:? 


402 


Index 


Observation,  method  of,  ii,   278 

Odors  of  animals  and  plants,  i,  84 

CEdema,   ii,   116 

Oil-drops  and  globules,  in  eggs,  i, 
213,  215 

Oliver,  J.  R.,  ii,  7 

Omnipotence  of  chromatin  in 
heredity,  ii,   14 

One-cell  stage  of  organism's  life, 
i,  214 

Ontogeny,  misuse  of  term,  i,  271 ; 
of  protozoa,  i,  277;  effort  to  ex- 
plain on  elementalistic  princi- 
ples, ii,  158 

Optic  nerve,  ii,  164 

Organ,    i,    245 

Organ-forming  substances,  in  the 
ovum,  ii,   16;  theory  of,  141 

Organ-germs,    i,    209 

Organelles,   i,   248 

Organic  formation,  and  isolated 
fragments,    i,    176 

Organic   matter,  i,   114 

Organicists,    i,    7 

Organ-independence,  i,  40 

Organism,  as  a  whole,  familiarity 
of  expression,  i,  1;  definition  of, 
18;  distrust  of  by  biologists,  25; 
and  its  chemistry,  75;  and  its 
protoplasm,  120;  and  its  cells, 
150;  in  interpreting  the  cell,  156; 
substitution  of,  for  cell,  194; 
consisting  of  one  cell,  227;  fic- 
tion of  structureless,  256;  as 
cause,  276;  "organless,"  232, 
320;  hypothetical  primitive,  319; 
fundamentally  dynamic,  ii,  134; 
"body"  and  "soul"  combined, 
151;  as  causal  explanation,  199; 
as  cause  of  chemical  transfor- 
mation, 205;  a  natural  object, 
207;  attributes  and  acts  of,  215; 
living,  275 ;  physico-chemical 
conception  of,  and  conscious- 
ness, 324 

Organism-transforming  action  of 
thyroid   substance,  i,   145 

Organismal  theory,  i,  2,  280;  con- 
structive side  of  discussion  of, 
ii,  91;  and  C.  M.  Child's  results, 
111;  and  elementalist  stand- 
point,   148;    in   relation    to   trop- 


isms,   188;  of  consciousness,  282 

Organismalism,  i,  2;  and  correla- 
tion, 17;  remarks  on  the  term, 
28 

Organismalist,    ii,    149 

Organization,  law  of  eml)ryonic 
development,  i,  14;  of  infusoria, 
282;  of  chromosomes,  ii,  28; 
physical,  and  instinct,  310;  and 
emotion,  316 

Organizing  power  of  living  beings, 
i,  211 

"Organless  organisms,"  i,  232,  320 

Organogenesis,  i,  325 

Organoids,  i,  248 

Organs,  fallacious  teachings  about, 
i,  242;  rudimentary,  277;  "can 
belong  only  to  multicellular  or- 
ganisms," 281 

Origin  of  species,  i,  4 

Osborn,  H.  F.,  i,  320 

Outer  layer,  universality  in  or- 
ganisms,  i,  301 

Ovum,  i,  210;  as  entity  and  as 
germinal  entity,  ii,  15 

Oxidation,  in  animal  body,  i,  340; 
ii,  346 

Oxygen,  "doscarecious"  powers  of, 
ii,  204;  in  relation  to  conscious- 
ness, 290  et  seq.;  as  respiratory 
substance,  301 ;  latent  attribute 
of,  341 ;  hereditary,  ontogenic 
and  individual,  347;  "activation" 
of,  348 

Pacific  Ocean,  iournev  of  fur 
seals  in,  ii,  211 

Pain,  ii,   183 

Pairing,  promiscuity  in,  ii,  267 

Paleobotanists,   ii,   58 

Paleontologists,  i,  322 

Palms,  ii,  299 

Pancreas  and  pancreatic  juice,  ii, 
120 

Pangens  of  Darwin,  i,   19 

Parallelism,  psycho-physical,  irre- 
solvable inconsistency  of,  with 
organismal  standpoint,  ii,  150, 
220;  historical  basis  of,  297 

Paramecium,  i,  326;   ii,  253 

Paratoid  gland,  of  toad,  i,  111 

Parthenogenetic   eggs,   i,  351 


Index 


403 


Particles,    ultimate,    ii,    1.>1 

Partisanship   in  science,  i,  338 

Pavne,   F.,   ii,  3i 

iviirl,    R.,   i,  311 

Pecklianis,    G.    W.    and    K.    S.,    ii, 

251 
Percepts,  ii,  291. 

Periodic  law,   in   chemistry,   ii,  32?) 
Personal    conscience,    ii,    292 
Personality,    ii,    33.>;    and    ])erson, 

295;   and  elementary   sid)stances, 


327 


Petrels,   i,  85 

Phases,  fluid,  of  cell  system,  i, 
216;   of  the   cell,  ii,  32() 

Philoso])her's   stone,  ii,  288 

Philosophy,   cartesian,  ii,   297 

Phleps,  Ed.,  ii,  135 

Phlofiiston,   i,   225 

Phos])horus,  ii,  290;  a  simple,  287; 
jrlow  of,  343 

Phylogeny,  of  biochemical  sub- 
stances,  i,   110 

Phylloxerans,    eggs   of,    i,    352 

Physical  basis  of  life  (Huxley),  i, 
121 ;  of  heredity,  ii,  64 

Physical  chemistry,  in  biology,  i, 
114;  and  protoplasm  doctrine, 
140;  and  the  organisnial  stand- 
point, 191;  conception  of  cell, 
215,  333;  conception  of  organ- 
ism, ii,  71 ;  limitations  of,  in 
biology,  208;  absence  of  in 
earlier  biology,  336;  anti-ele- 
mentalistic   tendency    of,   341 

Physical   science,  ii,   152 

Physio-chemical  substances  and 
forces,  in   ])ehavior,  ii,  312 

Physics,  and  chemistry  in  hered- 
ity, i,   115;  ])rovince  of,   141 

Physiologist,  ii,  285 

Physiology,  and  heredity  of  mus- 
cle fil)ers,  ii,  61  ;  distinctive  task 
of,  274 

Pbytin,   ii,   123 

Pigeons,  control  of  sex  in,  ii,  78; 
Whitman's  studv  of  habits  of, 
314 

Pigment,  from  chromatin,  i,  338; 
bearers,  liK) 

T*igmcntation,    i,    2V.\ 

Pill   bugs,  i,  219 


PiilslMiry,    W  .    n.,   ii,   231 

Pineal   body,   ii.    III 

Pituitary    I'iody,   ii,    1  I  1.    \!\ 

Plankton,    ii,    280 

IMantagens,   i,   36 

Plmutrkt,    ii,    109 

IMasma,   i,   134 

Plasmic    elciiicntalisni.    i,     ill 

Play    of   animals,    ii,    .'73 

Pleasure  ai\(l  Pain,  criterion  of,  i, 
289 

Pleoniorpbism,   i,   2(\ii 

Plover,   Clolden,   ii,   259 

Pluralism,    j)hilos(ipliic,il,    ii,    29t 

Pluleus   larva,  i,  203 

Poisoning,   strychnine,   ii,    173 

i'olaritv,  in  plants  and  animals,  i. 
181 

Pollen  grains,  in  fc-rtili/.ation,  I, 
34!i;  structure  of  mcmbraiu-  of, 
ii,   57 

Poronjfora  tjujantea,  i,  270 

Post-generation,   i,   207 

Postulate,    ii,    297 

Powers,  secret,  of  substances,  ii, 
300 

Precipitin   reaction,  i,    100 

Predisposition,   i,   206 

Primitive,   i,   290 

Primrose,  evening,  i,  'Mi\\ 

I'rinnni    morrn.s,   i,   276 

Priru-iple,  of  unity  of  organiza- 
tion,  i,    166;   of  aggri'gation,    182 

Principles,    Alcln'mists',    ii,    288 

Private  oi)inion,  ii,  292 

Promisiuity   in  mating,  ii,  2(»7 

Promorphology,  of  gi-rm  c«-lls.  i. 
211;   metaphysical,  225 

Prop<-rties  of  the  whole,  i,   13,   1  \^ 

Pro|)li\  siolog^•,  i,  212 

Protista,   i,    2'M),   280 

l*rotoj)hyta,   i.    23(» 

Protoj)lasm,  as  goal  of  l»iology,  i, 
5;  mystification  of,  12(>;  at\<l  tiM' 
organism,  120;  .Max  S<'hult/.e  on, 
1.'5;  latest  \  lews  MS  to  morphol- 
ogy of,  ][VA;  sjieciHcity  of,  113; 
as  general  term  mu-*  ''i-  used  iu 
plural   muuber,   I  Is 

Protozoa,  tlieor<'tical  simplicity  of, 
i.  J'.\(),  2S6;  (lr\rIopiu<'nt  of,  ?67 ; 
theoretical      aggn-galioii      of      to 


404 


Index 


produce  metazoa,  368 

Protozoan  colony,  theory  of,  as 
nature  of  metazoan,  i,  222 

Protozoology,  i,  280 

Pseudopodia,  i,  240;  pseudopodial 
system,   239 

Psychic  activities,  subrational, 
four  certainties   about,  ii,  250 

Psychic  integration,  ii,  94;  discus- 
sion of,  214  et  seq. 

Psychic  life,  subrational  moiety 
of,  ii,  246;  phases  of,  2T4;  spec- 
ificity of,  276;  man's  higher, 
283;  catholicity  of  attitude 
toward,  284 

Psychical,  organic  connection  be- 
tween  physical    and,   ii,   239 

Psychical  element,  an  abstraction, 
ii,  235 

Psycho-analysis,  ii,  350 

Psycho-physical  parallelism  (see 
parallelism) 

Psychoids,  ii,   149 

Psychologist,   ii,   285 

Psychology,  associationist,  ii,  228, 
and  Wm.  James,  229;  without  a 
soul  and  without  a  body,  322; 
social  and  domestic,  332 

Purpose,  of  reflex,  ii,  184 

Python,  proportionality  of  parts 
in  skeleton  of,  ii,  96 

Radicals,  compound,  in  chemistry, 

ii,  343 
Radiolarian,    compared   with   jelly 

fish,  i,  235;  swarm  spores  of,  278 
Rage,  ii,  183 

Random  movements,  ii,  252 
Reaction,  chemical  and  neural,  ii, 

286 
Reason,      ii,      216;      "forms      the 

world,"   243 
Receptors,    superficial    and    deep, 

ii,   173 
Redwood,   adventitious  buds   in,  i, 

38;  meristic  growth  series  in,  ii, 

101 
Reed,  H.  S.,  on  adventitious  buds 

in  lemon  trees,  i,  38 
Reflex  arc,  cellular  integration  in, 

ii,   163 
Reflexes,  simple,  an   abstract  con- 


ception, ii,  168;  compounding 
and  spreading  of,  171 ;  scratch, 
allied,  proprio-ceptive,  172;  an- 
tagonistic, 174,  176;  inhibition 
of,  and  compensatory,  177;  pur- 
pose of,  184;  relation  to  in- 
stinct,  188,   246 

Regulation,   formative,  i,  199 

Reichert,  E.  T.,  i,  95 

Relationship,  the  problem  of 
causal,  ii,  224 

Rennet,   i,    104 

Repetition,  in  organic  growth,  ii, 
95  et  seq.;  in  instinctive  activ- 
ity, 257  et  seq. 

Repetitive  parts,  ii,  95 

Reproduction,  asexual,  and  hered- 
ity, i,  309  et  seq. 

Researches  in  biology,  field,  labor- 
atory, quantitative,  ii,  278 

Resemblance,  importance  in  doc- 
trines of  heredity,  i,  312,  315; 
and  difference,  317 

Respiration,  life,  and  conscious- 
ness, ii,  286  et  seq.;  of  muscle, 
346 

Responses,  ii,  216 

"Restlessness,"  mental,  ii,  225,  243, 
307 

Retzius,  G.,  comparative  re- 
searches on  spermatozoa,  i,  216, 
ii,  2 

Reversibility,  of  direction  in  nerve 
conduction,  ii,   166 

Rhizopod,  i,   123 

Rhus,  chemistry  of,  i,  87 

Riddle,  O.,  ii,  76 

Ritter  and  Forsyth,  ii,  45 

Robertson,  T.  B.,  on  chemical  ac- 
tion in  growth,  ii,  105;  on 
tethelin,   123 

Rousseau,  J.  J.,  ii,  224 

Roux,  W.,  terminology  of  "De- 
velopmental mechanics,"  i,  18; 
Mosaic  theory  of,  198 

Royce,  J.,  relation  between  trop- 
isms  and  apperception,  i,  23,  ii, 
220;  on  mental  initiative  and 
restlessness,  243 

Ruzicka,  V.,  i,  262 

Sachs,  J.,  i,   16;   law  of,  219 


Index 


405 


"San    Diego    region"    (oceanic),   ii, 
281 

Saint-Hilairc,  J.,  i,  7 

S.i  jous,  K.  de  M.,  ii,  124 

SalaiiKindcr,   ii,  'ill 

Salmon,   ii,  2b0 

Salj)a,   i,   31(> 

Sarcode,  identification  with  j)lant 
])r()toj)lasin,   i,    \2\\ 

SchiifiT,  1',.,  on  endocrine  organ.s, 
ii.  It;  drng-like  action  of  inter- 
nal  .secretion,    122 

Scliandinn,   F.,   i,  201 

Sclileiden,  M.  J.,  conception  of 
plant,  i,  34 

vSchnltze,  Max,  on  protoplasm  and 
cell,  i,  125  et  seq. 

Sdiwann,  Th.,  theory  of  cells,  i, 
150 

Science,  positive,  1,  208;  creative 
impnlse  in,  ii,  227 

Scientific,  spirit,  i,  338;  attitude, 
difference  between,  and  philo- 
sophic, ii,  307 

Scratch-reflex,  workings  of,  ii, 
172;  possiljle  "ultimate  explan- 
ation"  of,  202 

Scripps  Institution,  statistical 
methods  in,  ii,  280;  for  Biolog- 
ical   Research,  332 

Sea  Urchin,  properties  of  eggs  of, 
i,  107;  experiments  on  develoj)- 
ing  eggs  of,  202 

"Seat"  of  heredity,  i,  321;  of  in- 
heritance material,  ii,  23;  brain 
supposed  seat  of  coordination, 
ii,  191 

"Secret  powers"  of  substances,  ii, 
341 

Secretin,  ii,  120 

Secretions,  internal,  integrative 
office   of,  ii,   113  et  seq. 

Secretory  systems,  internal,  ii,  128 

Sedgwick,  A.,  ii,  50 

Seeds,  gradations  in,  i,  102 

Segmental  theory  of  nerve  action, 
ii,    185;   of   nervous  system,   190 

Segregation,  in  heredity,  i,  355 

Selection,   sexual,    ii,   2(j2 

Self-})reservation  and  regulation, 
i,  18;  -differentiation,  199,  208; 
ii,   245;   -activity,   22G;   -exhaust- 


ing ceremonies,  2G4;  -injury 
through  M-x  imptdse,  2«i7;  nat- 
ural history  method  in  study  of, 
2H2;  -defenee  aiul  -preserviitioM, 
2\)2\  or  jH-rson,  3(»l ;  -<-ontrol, 
305;    -development,   305 

Sellars,  H.  W.,  indivichiality  in 
percepts,  ii,  2«>1;  the  iiulividuid, 
and    social    psychology,    'X\2 

Si-iisalionalism,   ii,   JIH 

Sense  organ,  ii,    l(»5;  senses,  ilfi 

t>equoi<i  nemperviretui,  adventi- 
tious buds  of,  i,  SS;  growth 
.series    in,    ii,    lui 

Series,  graded    n-petitive,   ii,  95 

Serum,   rabi)it,   i,    101 

Sex,  cell  production,  theory  of,  i, 
<)1;  as  unit-character,  3tS;  de- 
pendence upon  chromosomes, 
34(),  350;  impulse,  excessiveness 
of,    ii,   265;    power   of,   2(i7 

Sexes,  numerical  j)roportion  of,  ii, 
2(J7 

Shakesj)eare,  Win.,  "rtrkless  volu- 
bility" of,   ii,  223 

Sherrington,  S.  C,  fundanu-ntal 
work  on  nervous  system,  i,  22, 
ii,  1G2  et  seq.;  lu'glect  of.  by 
J.   Loeb,   185 

Shrikes,  ii,  270 

Simj)l(;  reflex,  an  abstract  coiu'ep- 
tion,   ii,   1()8 

Simi)licity,   "ultimate,"   i,   3-'() 

Skin,  universal  j)resence  of,  in  or- 
ganisms, i,  300 

Smallpox,   i,   2(»4 

Smell,   i,  84 

Smith,   J.    H.,   i,   211 

Society,    and    individual,    ii,    'X\2 

Soma,  i,  319 

Song    habits    of    birds,    ii,    260 

Soul,  com])osed  of  "sen)iiud 
atoms,"  i,  3;  as  aspect  of  organ- 
ism, ii,  215;  aiul  botly,  interac- 
tion   bet\v«'en,   \\2".\ 

S|)ecialist,  the   unpoiseil,   ii,   247 

Species,  difference  in  egg  of,  ii,  20 

Specific  tlifferences  In-tween  germ 
cells,   imj)ortance  of,   i,   2\i 

Specification  of  orL'^anic  matter,  i, 
111 

Specificity,     clu'mical,     of     organ- 


406 


Index 


isms,  i,  83;  of  corresponding 
proteins,  95;  of  protoplasms, 
143;  of  sperm  of  anthropoids 
and  man,  ii,  2;  of  animal  be- 
havior, 276;  of  psychical  and 
reactive  life,   281 

Speculations,  i,  393 

Spermatozoa,  species  differences 
in,  i,  216;  structure  of  tail  of, 
333;  structure  of  head  of,  342, 
and  ii,  87;  two  kinds  of,  i,  347; 
subject  to  heredity,  399;  resem- 
blance to  tadpole,  ii,  3;  ontog- 
eny  of   mammalian,   4 

Spicules  of  sponges,  ii,  50 

Spinules   of   Ascidian,   ii,  44 

Spirits,  historical  relation  to  es- 
sences, ii,  288;  historical  relation 
to  breath   and  air,  338 

Sponge   tissues,  i,   144 

Spontaneous,  generation,  ii,  316; 
in  mental  activity,  226;  on 
meaning  as  applied  to  "origin 
of  life"    (see   glossary) 

Spores,  i,  269;  of  mosses  and 
ferns,  ii,  258 

Squid,   i,  221 

Squirrel,  storing  habits  of,  ii,  271 

Stage,  one  celled,  of  animal  life, 
i,   214 

Star  fishes,  ii,  96 

Starch  grains,  i,  214 

Stein,  Fr.,  i,  286 

Stentor,  supposed  nerves  in,  i,  245; 
ontogeny  of,   272 

Stephens,  Frank,  on  storing  habits 
of  bees,  ii,  268;  of  antelope 
ground   squirrel,   271 

Stevens,  Miss  M.  M.,  i,  350 

Stimuli,  summation  of,  ii,  167 

Storing  habits,  of  honey  bee,  ii, 
268;  of  woodpecker,  269;  of 
mammals,    270 

Stout,  G.  F.,  ii,  298;  unanalyzed 
cognition  in  consciousness,  308; 
on  personality  in  animals,  328 

Strassburger,  Ed.,  i,  342;  and 
chromosome   dogma,   ii,   259 

Striated  muscle  tissue  and  hered- 
ity, ii,  59 

Structurelessness,   i,   285 

Structures,    ii,    199;    constitutively 


antagonistic,     134;     organ-form- 
ing,  141 

Struggle,  of  the  parts,  ii,  175;  for 
existence,    225 

Studies,  practical,  i,  214 

Sub-conscious,   ii,  350 

Subjective,  ii,  231;  side  of  asso- 
ciation,  230;   and   "inner,"   292 

Substance,  of  soul,  i,  4;  organic 
production  of,  81 ;  simple  homo- 
geneous, 281;  imaginary,  240; 
nutritive,  244;  criterion  of  ele- 
mentary, 287;  abuse  of  the 
term,  298;  respiratory,  301;  and 
energies,  337;  free  and  vague 
appeals  to,  ii,  239 

Sugar,  and  enzyme  action,  ii,  81 ; 
from   liver   into   blood,   132 

Summation,  of  stimuli,  ii,  167 

Sumner,  F.  B.,  ii,  258 

Supernaturalism,  and  finalism,  ii, 
142;  and  materialism,  148 

Suprarenal  body,  ii,  114,  135;  in 
alliance  with  thvroid  and  pitui- 
tary,   127 

Surface  energy,  of  muscle  fibers, 
ii,  63 

Surfaces   of  separation,   ii,    170 

Surf  perches,  matings,  habits  of, 
ii,  265 

Surgeons,   Swiss,  ii,  115 

Surplus  energy,  ii,  273 

Sutton,  W.   S.;  i,  356 

Swarm  spores,  of  radiolaria,  i,  278 

Swezy,  olive,  i,  291 

Symbiotic  theory  of  organism,  i, 
'35,   183 

Symmetry  of  animals,  ii,  189 

Sympathetic  nervous  system,  ii, 
"128 

Synthesis,  organizing,  ii,  149;  as- 
similative, 205;  and  analysis, 
235;  apperceptive,  237 

System  of  nature,  i,  4 

Systematic    biochemistry,    i,    95 

Tadpoles,  of   frogs   and   toads,  ii, 

143 
Takamine,   J.,   ii,   123 
Taste,  i,  84 
Taxonomic,    in    organic    grades,    ii, 

42;  discrimination,  62 


Judex 


K): 


Taxonoinist,    i,   213 

Terms,  of  oxygen,  ii,  202;  of  life. 
207 

'IVtaiiv,  i,  117,  135 

'rclliciiii,  ii,  123 

TlK'ory,  of  struclive  of  jn-oto- 
})]asm,  i,  138;  wrong  Ix-ttcr  than 
none,  292;  Leil)nitzian.  ii,  1.50; 
of  organisms,  and  of  knowledge, 
152;  elemenlalist,  157;  iropistic 
and  segmental,  of  nerve  action, 
185;  of  tropisms,  232;  of  aj)per- 
eeption,  232;  of  knowledge,  29(); 
of   consciousness,   2})(> 

Tliomson,  J.  A.,  i,  30}) 

Threshold  of  excitability,  ii,   1<>5 

'I'hynuis,    ii.    111 

Thyroid,  ii,  121;  aj)])aratus,  111-; 
effect  of  removing,  115,  117; 
trij)le  alliance  with  pituitary 
and  adrenal,  127;  sul)stanc<-, 
143;  organ-transforming  sul)- 
stance   from,   145 

Tissue,  laboratories,  i,  s:i;  mix- 
tures, 143;  cultures,  1()8;  falla- 
cious teaching  about,  240;  of 
multicellular  organisms,  28 1; 
isolated,  294;  of  trees,  specific- 
ity of,  ii,   58 

Toads,   ii,    76 

Tonniges,  C,  i,  32(» 

Torrey,   H.   B.,  ii,  97 

T<)ti|)otence,   theory   of,   i,   202 

Transformation,  essence  of  evolu- 
tion, i,  41 ;  hereditv  works  by,  i, 
312,  322;  of  substances,  ii,  72, 
287 

Transmission,  in  ccumection  with 
heredity,  i,  312 

Transmutation  of  metals,  ii,  288 

Transcendentalism,    ii,    233 

Treviranus,  i,  5 

Trial  and  error,   ii,  2>>2,  2M) 

.Trichocysts,  i,  326 

'i'rij)ylea,  i,  278 

'i'ropism  theory,  essentially  an  or- 
ganismal  theory,  ii,  188;  relation 
to    apj)erce})tion,    232 

Tropisms,  explained  l)\  organism, 
ii,  190;  organismal  nature  of, 
239;  automatic  and  anticipatory 
character     of,     241  ;     higher     ra- 


tional   life   and,      ' 
Tropistic    and    higiiri     p^vmu      «<  - 

livity,  ii,  22U 
'iropistic    mechanism,    ii.    !'''• 
Trojjistic   tlK'ories  of  nrrvr   nrtion. 

ii,    IH5 
Truth,    idtimatr,    ii.    i;,.».    >89 
Trypsin,    i,    loii 
I'ype,   i,    7 
Tyranny,    of    the    whole,    ii.    15H 

ritimatr,   proi»lini,   i,  35;   criterioi 
of,  ii,    !  W»,   JO!  ;   |)artieles,  ii,   151, 
truth,     152.     175,     2H9;    elements. 
228 

ntramicroscopic  organisms,   i,  .»«»5 

Inic-ellular,  i,  290 

I'nits,    representative,    i,    30(j 

I'nity,  tlu"  organism  the  "only 
real,"  i,  1 ,»,  2i\,  205;  physiolog- 
i<al,  11;  of  th<-  indi\idual,  'X\; 
thought   of,   ii,   150 

I'ranium   H.,  ii,  311 

rtility,  racial,  ii,  251;  natural  se- 
lectionist   meaning   of,    261 

\'ariation,    ii,    245 

N'ertebrates,  man  a,  because  moth- 
ers  were,  ii,  43 

\'erworn,    M.,   ii,    J7f> 

N'etch,    ii,    IO<i 

N'irgin    propagation,    i,   352 

N'ital    force,  ii,   I  1!) 

N'italism,  i,  113;  author's  attitncK* 
toward,   ii,   207 

\iviparous  bony  fishes,  mating 
habits  of,  ii,  2<»5 

N'olition,    ii,    161 

Wallace,    A.    |{.,    ii.   278 

Waller,    H.    K.,   ii,    126 

Warblers,    ii,   264 

Wasps,  instincts,  variidiility   of,  ii. 

251;   excessive   action   «»f,   2(»8 
Wassermann,  i,  258 
Watasc,    S..    i.   221 
WCbber,    11.   .1.,   i,  :W() 
Weismann,   A.,  studies  <»f  srx-<TlU 

in    hydroids,    i,    (»<»;    («r!tr's    rv- 

sults   i*ontrar\    to.   JiS;   inrtapln  s- 

ics  of,   2.V.,  3  IS 
W  hale,    breaching    of,    ii,    257 
\\  heeUr,     W       M.    wiisr    of    snirll. 


408 


Index 


and  odors,  of  ants,  i,  87,  89; 
early  embryology  of  insects, 
218;  as  field  naturalist  of  mod- 
ern type,  ii,  278;  on  problem  of 
instinct,  284;  on  instinct  and 
bodily   organization,   311 

Whitman,  C.  O.,  as  pre-organis- 
malist,  i,  11;  on  cell-theory,  220; 
relation  between  instinct  and 
structure,   314 

Whole,  embryo,  i,  204;  "tyrannizes 
over  parts,"  ii,  159 

Will,  feeling,  and  intellect,  ii,  216, 
217 

Wilson,  E.  B.,  as  pre-organismal- 
ist,  i,  11;  on  cell  structure,  135; 
statement  of  cell-theory,  151 ;  on 
"real  unity,"  192;  on  early  em- 
bryology of  amphioxus,  204;  on 
promorphologj^  217;  on  x  and  y 
chromosomes,  350;  on  connec- 
tion between  chromosomes  and 
Mendelian  inheritance,  356;  pro- 
posal to  drop  "determiner"  as 
genetic  term,  ii,  82;  on  germ  as 
detached   portion   of   parent,  88 


Wilson,   H.   v.,  i,   144 

Winterstein,   H.,  ii,   62 

Woodpecker,  California,  storing 
habits  of,  ii,  269 

Wood-tissues,  ii,  58 

Work,  energy,  power,  force,  ii,  342 

World,  external,  ii,  303 

Wundt,  W.,  and  apperception,  i, 
23;  definition  of  apperception, 
ii,  233;  seeming  transcendental 
element  in  the  apperception  of, 
234 

Yerkes,  R.  M.,  on  combined  ex- 
perimental and  field  research  in 
behavior,  ii,  279 

Ziegler,  C,  i,  201 
Zoja,  R.,  i,  204 
Zoological  diagnosis,  i,  265 
Zoologist,  anthropological,  ii,  285 
Zoology,  instruction  in  elementary, 
i,    236;    and    the    science   of   be- 
havior,     ii,     208;      fundamental 
terms  of,  247;  taxonomic,  276 
Zygote,  i,  269 


